Strange Natures: Conservation in the Era of Synthetic Biology 9780300258677

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Strange Natures

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Strange Natures Conservation in the Era of Synthetic Biology Kent H. Redford and William M. Adams

New Haven & London

Published with assistance from the foundation established in memory of Amasa Stone Mather of the Class of 1907, Yale College. Copyright © 2021 by Kent H. Redford and William M. Adams. All rights reserved. This book may not be reproduced, in whole or in part, including illustrations, in any form (beyond that copying permitted by Sections 107 and 108 of the U.S. Copyright Law and except by reviewers for the public press), without written permission from the publishers. Artwork by Emily Elsner. Yale University Press books may be purchased in quantity for educational, business, or promotional use. For information, please e-mail [email protected] (U.S. office) or [email protected] (U.K. office). Set in Galliard Old Style and Gotham types by Integrated Publishing Solutions. Printed in the United States of America. Library of Congress Control Number: 2020947561 ISBN 978-0-300-23097-0 (hardcover : alk. paper) A catalogue record for this book is available from the British Library. This paper meets the requirements of ANSI/NISO Z39.48–1992 (Permanence of Paper). 10 9 8 7 6 5 4 3 2 1

Dedicated to Pamela Shaw and Francine Hughes

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Contents

Preface  ix Acknowledgments  xiii 1 The Place of Nature  1 2 The Problem of Nature  20 3 Nature’s Diversity  45 4 Conserving the Genetic Pieces  68 5 Rewiring Nature  89 6 Synthesizing the World  107 7 Genetic Technologies in Conservation  136 8 Nature’s Future  167 9 Conserving Strange Natures  191 Appendix: Scientific Names of Species  213 Notes  221 Index  265

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Preface

Nature is becoming strange. Its species and ecosystems, penguins and meerkats, and woodlands and savannas are familiar from children’s books and television documentaries. For more than a century, conservationists have worked to protect this natural world. Yet its diversity, which evolved so gradually, is being lost with disconcerting speed. Human demands on the biosphere expand seemingly without limit, and their impacts affect all parts of the earth. In this world, new and strange natures are being called into being. Ecosystems have always shifted and changed as they are worked and reworked by the forces of climate, geology, and evolution. Now nature is increasingly having to adapt to the new conditions created by human action. But the strangeness of nature goes beyond this enforced hybridity, and the mark of human hands is growing stronger. Scientists have now developed the ability to edit the genes of living species and to create novel forms of life. This science, synthetic biology, allows nature to be reshaped more or less at will. The power of technologies to engineer nature at the ecosystem scale may be familiar, and conservation strategies to limit change are well developed. The possibility of engineering genomes is new—and strange. Synthetic biology is based on the development of tools that can be applied at the genetic scale, cutting, pasting, and rewriting individual components of DNA. These tools offer the power to shape living cells and organisms in unprecedented ways. Gene-editing tools such as CRISPR ix

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are already transforming biotechnology and agriculture, as well as human medicine. Genetically engineered algae growing in industrial vats are yielding biological products once extracted from plants or other species. Crops are being engineered to grow faster or to resist pests, and pest species themselves are being engineered to be less effective. Synthetic biology is expected to drive dramatic changes in economies and societies both as revolutionary new biological production systems take root. Such powers to rewrite the genomes of living organisms offer both threat and opportunity to nature. On the one hand, there is the potential risk to biodiversity of engineered organisms being released into the wild and affecting species and ecosystems. On the other hand, the same technologies offer previously unimagined possibilities as tools for conservationists themselves. There is discussion of the prevention of wildlife diseases that threaten the extinction of rare species, or the control of invasive species on oceanic islands. There is even discussion of bringing extinct species back from the dead. Gene editing makes strange natures, but, perhaps most problematically, it makes the idea of nature itself strange. Nature is usually imagined as separate from the domain of people. We think of the natural world as beautiful, diverse, valuable, and “natural,” whereas the human world is developed, artificial, and increasingly unnatural. Gene editing erodes the difference between what is natural and what is human-made. In doing so it threatens to destabilize the very idea of nature that provides the foundation for conservation. This book is about the challenge of genome editing for our idea of nature and our ideas about its conservation. It has been about ten years in the making. We met in 2004, when Bill was invited to join the team planning a project funded by the MacArthur Foundation, “Advancing Conservation in a Social Context,” of which Kent was already a member. Meetings in Switzerland, Chicago, Madagascar, and Athens, Georgia, revealed a shared obsession with the curiously varied ideas people have about nature and how to conserve it. Our backgrounds are very different. Bill was trained as a geographer and has worked primarily in academia, writing about the history of conservation and sustainability and the clash between conservation and development. He did his doctoral research on the downstream impacts of



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dam projects in Africa. Kent is an ecologist and did his doctoral work on giant anteaters in Brazil. He has worked both in university research and in the professional conservation sector, with the Nature Conservancy and the Wildlife Conservation Society. We share a passion for the natural world and a fascination with the different ways people engage with it. We have lived our adult lives watching the erosion of living diversity, at home in the United States and the United Kingdom, and internationally. We have read libraries full of books and journal papers and contributed to more of them than we care to remember. We have analyzed the past and tried to predict the future; we have lamented and preached, celebrated and complained. We have tried to question entrenched thinking about conservation, both inside and outside the movement. And all the while, we have watched biodiversity continue to leak away, destroyed by the very economic engine that creates human wealth and welfare. Conservation has never been so challenging—or so important. In 2013 we worked with the late Georgina Mace, ecologist and conservationist, to organize a meeting of engineers, biologists, and conservation scientists and professionals to explore the implications of synthetic biology for the conservation of nature. The meeting took place over two days in Cambridge in the United Kingdom, bringing together enthusiasts and skeptics and promoters and opponents of the new technologies to start thinking about the opportunities and risks of new genetic technologies for conservation. Since that meeting, we have gone on working on conservation dimensions of emerging genetic technologies and especially their implications for the interlaced questions of naturalness and artificiality. The rise of novel genetic technologies is by no means the only potential future threat to biodiversity, and it is certainly not yet the greatest. But it poses fundamental questions about how conservation should be done in a world of strange natures. Artificial techniques such as fencing and burning are already widely accepted in conservation management. Should synthetic biology tools be thought of in the same way? Or is there something different about their strangeness and unfamiliarity? Thinking about this requires an understanding of genes and evolution, of how the technologies of genetic engineer-

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ing and especially genome-editing work, of the ways these technologies are already being applied in biotechnology and agriculture, and the ideas being discussed for using them in conservation. It also demands that we think about how best to have oversight of innovations in synthetic biology and how to address concerns about risks. Above all, it requires understanding how the capacity to edit genomes of wild species changes our thinking about nature. This book has been a challenging experiment in collaborative writing. We come from different intellectual backgrounds, formed by different geographies, experiences, and traditions. We have had to address areas of science and scholarship in which we were never trained. We have had to struggle to understand how disparate areas of knowledge fit together and to think through how we each (and together) respond to key questions. We have debated with each other over computer screens, pub tables, carpets of bluebells in ancient woodlands, and the sites of dam removals, where sea-run alewives course up rivers that had been blocked to them for centuries. We have done our writing on opposite sides of the Atlantic, mastering the dark arts of electronic communication to share our thinking, concerns, and emotions as best we can. We have enjoyed navigating the backwaters of American and British English and finding a common language and style. The growing powers conferred by synthetic biology reflect the ever-­ extending human transformation of the biosphere. The capacity to reorder DNA, genes, and genomes to fit human plans challenges our notions of humanity’s place in nature. They are reminders of the importance of genes and genetic engineering to human futures, as well as to the future of conservation. Bill Adams, Cambridge, United Kingdom Kent Redford, Portland, Maine, United States April 2020

Acknowledgments

We owe a huge debt to colleagues and friends for their assistance with this book. Over the years we have had too many conversations with too many knowledgeable people about the things we discuss in this book to thank everyone individually. It goes without saying that as squirrels of the conservation and research worlds, we have been snapping up unconsidered trifles for years, not always under conditions where we could take notes. First and foremost, we would like to thank Lisa Adams of the Garamond Agency for having faith in us, and Joe Calamia at Yale University Press for taking us on. We are very grateful to Jean Thomson Black, Elizabeth Sylvia, and the team at Yale University Press for all their work on the manuscript. We thank Emily Elsner for her wonderful drawings that are included on the chapter-opening pages. Bill is grateful to all the research students and postdoc students he has worked with over the years for the education they have so willingly provided. He would particularly like to thank Maan Barua, Matthew Gandy, Phil Howell, Chris Sandbrook, Adam Searle, Sverker Sorlin, and Jonny Turnbull for their advice and support while this book was written. Over the years the Department of Geography’s Political Ecology Group and Vital Geographies Group, as well as the Synthetic Biology Strategic Research Initiative and the Conservation Research Institute at the University of Cambridge have kept him on his toes and taught him a great deal. Kent would like to thank Steve Sanderson for believing in the importance of this idea and supporting the Cambridge meeting in 2013. He xiii

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would also like to thank colleagues at the various meetings on synthetic biology and conservation, in particular Ryan Phelan and Claudio Campagna. The International Union for Conservation of Nature (IUCN) Task Force and its Technical Subgroup provided a rich community of people struggling to find honest ways through a complicated and contentious topic. He would particularly like to thank Tom Brooks and Nicholas Mac­farlane for their support; and for particular inspiration and support, Jonathan Adams, Elizabeth Bennett, Jason Delborne, Drew Endy, Todd Kuiken, Aroha Mead, Simon Stuart, and Delphine Thizy. In addition, key support and inspiration from Rob Carlson, Fred Gould, Ron Kreisman, Elaine Leslie, and particularly Craig Groves. We are enormously grateful to friends who read various chapters of the book: Lisa Adams, Christina Agapakis, Joe Calamia, Rob Doubleday, Nigel Dudley, Emily Elsner, Matthew Gandy, Fred Gould, Francine Hughes, Todd Kuiken, Chris Sandbrook, Pamela Shaw, Sharon Strauss, Barbara Thorne, and three anonymous referees. We greatly value your advice and have taken most of it. All remaining errors and infelicities are of our making alone. Kent could not have written this book or even gotten to this stage in his life without Pamela, who has believed in him, supported him. She has proved patient beyond words in encouragement, love, and an occasional kick. Thank you; this book is ours! Bill, likewise, owes a huge and continuing debt to Franc, who has endured his abstraction and repeated absences hunched over a laptop with remarkably good grace, and has been a loving, kind, knowledgeable, and insightful reader of drafts. Thank you, for this and previous book-shaped impositions on our lives. For both of us, our children—for Kent, Sofia and Hugh, and for Bill, Emily and Tom plus Emily’s new generation—are a constant reminder of the importance of the future and the state of the natural world within which we live our lives.

1 The Place of Nature

Come the hot days of summer, in late July and August, a small dark butterfly appears high in the canopy of oak trees in the woods of eastern England, its wings flashing purple and silver. The purple hairstreak butterfly, once common, is entirely dependent on oak trees and, hence, on the woodland that survives in small patches in an otherwise agricultural landscape. With it in the canopy move small migratory warblers, the chiffchaff and blackcap, and far below in the understory, the fallow deer and the badger. In spring, the woodland floor is bright with flowers characteristic of old woodlands—oxlip, wild garlic, and wood anemone. In May, the dark leaf mold is in places covered with a carpet of bluebells, flowering early before the canopy of trees closes over. These flowers are no recent arrivals. Bluebells were mentioned in Ray’s Flora of Cambridgeshire as long ago as 1660, and the Victorian English poet Gerard Manley Hopkins described their effect brilliantly, saying that the bluebell made “wood banks and brakes wash wet like lakes.” They still do.1 The contrast between the inside and the outside of these woodlands, in the landscape near to Bill’s home in Cambridgeshire, is stark. Inside, the ecosystem is brilliantly alive with diversity and life. The bright summer light penetrates a few meters only and is then lost. Outside, an endless sky

Purple hairstreak butterfly 1

2 The Place of Nature

stretches over a gently rolling landscape of seemingly perfect uniformity. This is good arable farming country, and huge fields stretch as far as the eye can see, with barely a tree or a scraggy remnant of a hedge to break the smooth quilt of the growing wheat, barley, or canola (oilseed rape). The cultivation regime is intense, with mechanical applications of synthetic fertilizers, fungicides, insecticides, and slug and weed killers. Not many wild animals or plants survive the high-production management regime. Cambridgeshire’s woodlands are oases of biodiversity in a world of industrial farmland. Once, a thousand years ago, the woods were continuous across this landscape. Iron Age and Roman farmers cut them to make fields, as did the Saxons and other settlers. By a thousand years ago, the landscape was already agricultural. Nonetheless, these woods have been here for a long time. They contain elements of the fauna and flora that became established at the end of the last Ice Age, twelve thousand years ago. Many are surrounded by deep ditches and banks dug in the medieval period to mark and protect the woodland from livestock and people. They survived because they were too wet to cultivate, and because their owners valued the timber and wood products they provided. Hayley Wood west of Cambridge, for example, was named in a survey of the estates belonging to the Bishop of Ely in 1251. Its boundaries were more or less unchanged for the last seven and a half centuries, and for most of that time it was managed intensively to produce timber, poles, hurdles, and firewood.2 The conservation value of the small fragments of woodland that have endured in the lowland landscapes of the United Kingdom, and other industrialized countries, has long been recognized by conservationists. British ecologists call woodlands that can be shown from archival maps to have been continuously wooded for at least four centuries “ancient woodlands.” They not only hold species thought to have been characteristic of woodlands of previous millennia—in what ecologist Oliver Rackham called “the wildwood”—but they have very significant levels of biological diversity.3 Today’s “ancient woodlands” have been at the heart of British conservation strategies since the dawn of the conservation movement in the late nineteenth century. When a set of gentlemen naturalists founded the Society for the Promotion of Nature Reserves in London in 1912, ancient



The Place of Nature 3

woodlands were high on their list of priorities. The society’s foundation was a response to the ecological changes in the British countryside and its wildlife wrought by the Industrial and Agricultural Revolutions. Its stated aim was “to preserve from destruction in this country as much and as many as possible of the invaluable surviving haunts of nature.” These places were described as “the precious living relics of the world as it was, before man destroyed it.”4 Here is romantic nostalgia wrapped around a philosophy that nature was precious, was threatened, and could be (and indeed must be) kept safe from destructive human activities: human action saving nature from other human actions. Woodlands, grasslands, heaths, and moorlands were all surveyed, and a list prepared of nature’s most valuable “haunts.” There was irony in the fact that the society’s passion for “nature” was focused on ecosystems (woodlands, grasslands, heaths, and moorlands) that had all been shaped by human action over preceding centuries. The nature threatened by human action was already partly human-made, a tension that was bred in the bone of the modern conservation movement. It took a while to achieve the vision of nature reserves in the United Kingdom, but eventually it came to pass. By the end of the Second World War, the country had a conservation movement and the beginnings of a system of government designed to deliver conservation of nature that is still more or less recognizable today. Hayley Wood itself was eventually turned into a reserve in 1962, and conservation organizations now own many ancient woodlands across the United Kingdom. Many are designated by the government as Sites of Special Scientific Interest because of the species they support, and many are National Nature Reserves, the premier UK conservation designation.5 This model of conservation, based on the reservation of land for nature, now has counterparts all over the world. The establishment of systems of protected areas, from the wilds of Alaska to the rain forests of Kalimantan, is arguably conservation’s single greatest achievement. Every country in the world has some kind of conservation movement and has protected areas of land (and increasingly of sea) from development. In many countries, diverse ecosystems and rare species only survive because of this protection. Protected areas have become icons for the conservation movement,

4 The Place of Nature

from the first US national parks, like Yosemite or Yellowstone, to the savannas of Serengeti, Selous, or Maasai Mara, set aside in Africa after the Second World War as developing countries shook off colonial rule. By 2018, almost 15 percent of the earth’s land was in some kind of protected area. Many are large and draw the lion’s share of global conservation attention. Yet conservation work is not limited to them. In many countries, conservation efforts are also focused on smaller areas, like the British ancient woodlands, that maintain diverse species and ecosystems. Other examples include the thirty-five thousand or so remnant church forests of Ethiopia or the surviving fragments of Brazil’s Atlantic Forest (75 percent of which has been destroyed), now maintained in a network of small private, state, and national reserves.6 More than a century of formal nature conservation has therefore been built around the idea of nature as something separate from humanity that needs protection from human demands. At all scales, from the local to the global, successive generations of conservationists have labored to get reserves set aside to protect nature and to keep threatening humanity at bay. But how does this strategy work in the twenty-first century, when beleaguered nature seems to be everywhere in retreat in the face of climate change and continuing human expansion? Do we need to reconsider what we are saving? Are our tools fit for the purpose of saving the nature people care for? Or do we need to consider adding to our toolbox new ways that match the emerging understanding of the threats facing the natural world and the nature we wish to conserve? On a winter night in Cambridgeshire, Bill can still walk out into the dark. It is still (just) possible to get far enough away from streetlights, car headlights, and the ubiquitous floodlit world of roadside junk-food emporia to see the night sky. If you pick a moonless evening to stand and look up, maybe close to one of the surviving patches of ancient woodland, you see, of course, the stars, sometimes the soft illumination of the Milky Way. And every few minutes you see the blinking lights of jets setting off from one of the London airports for North America. If you keep looking, you can sometimes pick out a fainter zip of light, far higher up, moving at astonishing speed. This is the International Space Station, orbiting four hundred kilometers above Earth.



The Place of Nature 5

Looking down from planetary orbit, satellites scan a scrolling view of human occupation of Earth. This vantage gives the classic picture of Earth at night, a luminous darkness pinpricked and smeared with artificial light. It traces the tentacles of human occupation around the world, dense and bright in eastern China and the eastern and western seaboards of the United States and in northwestern Europe. In many other places the image is darker, either because population densities are low or lights are lacking. This is a map of industry, technology, and poverty—a chart of fossil fuels burning. The first satellites appeared in the night skies in the late 1950s, at the height of the Cold War. The Soviet Union was first, launching Sputnik 1 at the beginning of October 1957. It was tiny (just fifty-eight centimeters across) and orbited Earth every ninety-eight minutes until January 1958. Sputnik was a product of the Cold War. The “Space Race” it triggered led to profound changes in the way people thought about the planet. It made ordinary people come to terms, perhaps for the first time, with the fact that they were on a planet at all. The idea arose in the 1960s that Earth itself was no more than a spaceship adrift in the vastness of space. This idea is commonplace today, but it was first used by Adlai Stevenson, US ambassador to the United Nations, who said in 1965, “We travel together, passengers on a little space ship, dependent on its vulnerable reserves of air and soil; all committed for our safety to its security and peace; preserved from annihilation only by the care, the work, and, I will say, the love we give our fragile craft.” Through the 1960s and 1970s, the idea of Earth as a spaceship formed a powerful metaphor for global environmental concern, epitomized by the title of Buckminster Fuller’s Operating Manual for Spaceship Earth in 1968. The new environmental movement found defining icons in the Earthrise photograph taken by astronaut Bill Anders of Apollo 8 on December 24, 1968, showing Earth as a blue ball rising above the desolate surface of the lifeless moon, and the famous Blue Planet image of the whole Earth taken from Apollo 17 in 1972.7 The notion of the earth as a fragile and limited entity flew in the face of centuries of thinking about the planet as an endless and boundless resource for humankind. It had once seemed that fish filled the oceans, forests offered boundless supplies of timber, and unoccupied land offered free space

6 The Place of Nature

for agriculture. As the resources of one area became exhausted, imperial adventurers would go and annex another, claiming “new worlds” from people long settled there. Against this aggressive cornucopian model, the view from space offered a very different vision. It inspired the environmental movement’s concerns about limits to growth and human vulnerability. By the twentieth century, this argument was also centuries old, but it was made new in what came to be called the “age of ecology.” The new idea of nature was driven by two linked ideas. The first was about the limits to nature existing on a single planet in the vast lifeless distances of space. The second was of nature increasingly restricted on that planet itself, confined to protected areas and untransformed lands, ringed around by cities, industries, roads, and agricultural landscapes. These two ideas provided the root of today’s ideas about human responsibility for nature.8 Space exploration also provided data to shape this new ecological view of the earth and the living nature upon it. From the early 1970s, earth observation satellites started to stream back images of the earth’s surface. Unlike aerial photographs, their sensors covered everything, from poles to tropics, and recorded digital images that picked up different wavelengths of reflected light. Suddenly deserts and forests, cities and cultivated lands could be classified, mapped, and monitored as satellites passed repeatedly overhead. When viewed as a single tessellated whole, the earth was revealed to be in the process of radical transformation by human action, reshaped by technology and economy, restructured by patterns of human production and consumption. The newly imagined global nature was one pictured as under threat. In 2008, two North American geographers, Erle Ellis and Navin Ramankutty proposed a new way to classify the terrestrial biosphere to reflect the ecological patterns produced by human action. The conventional way to classify the distribution of ecosystems had been in terms of “biomes,” based on broad differences in vegetation types associated with variations in climatic patterns (desert, savanna, dry and moist forests, and so on). Ellis and Ramankutty took a different approach, analyzing the terrestrial biosphere in its contemporary, human-altered form. They mapped what they called “anthropogenic biomes,” or “anthromes.” Ellis and Ra-



The Place of Nature 7

mankutty’s analysis shows a world of ecological mosaics—a bewilderingly complex spatial blend of cities, towns, villages, croplands, residential rangelands, forests, and ancient woodlands.9 The conversion of the biosphere to human purposes has been gradual but accelerating. In 1700, nearly half of the terrestrial biosphere lacked human settlements or significantly transformed land use, and most of the rest (45 percent) was seminatural, with limited use for agriculture or settlement. By 2000, the opposite was true: less than 20 percent was seminatural and only a quarter undisturbed. The critical transition from mostly undisturbed to mostly anthropogenic ecosystems came early in the twentieth century.10 So, where is nature on this earth of human-transformed ecosystems? As the human footprint extended, nature found a place, or rather a set of places, remote from the hurly-burly of human ambition and hunger. James Watson, of the University of Queensland, and colleagues mapped what they called “the last of the wild,” by which they meant ecosystems showing no sign of built environments, crops, pasture, nighttime lights, railways, major roads, and navigable waterways on satellite images, and no other sign of high human population densities. The study calculated that 77 percent of land had been modified by human activities (excluding Antarctica, the only continent lacking permanent habitation), plus 87 percent of the ocean (influenced by pressure such as fishing, industrial shipping, and fertilizer runoff).11 The authors drew attention to the conservation value of these last wild areas for their biodiversity, ecosystem services, and as buffers against climate change. People do live in such areas and are often critical stakeholders in their management. For example, Indigenous peoples have tenure rights to over 40 percent of the area of terrestrial protected areas or ecologically intact landscapes. Yet it is here that the greatest numbers of species exist and survive and that ecosystems still function with minimal input from human industrial economies.12 The preservation of the integrity of ecosystems that are free from significant anthropogenic degradation is generally accepted as the most urgent priority for global conservation (although the lesser values of impacted ecosystems are also recognized). So, for example, “intact” forests

8 The Place of Nature

(those that have not suffered from fragmentation, logging, overharvesting, or changes in fire or flooding regimes) should be given priority in policy, planning, and conservation implementation.13 Despite the scale of conservation efforts, the loss of biodiversity—the extinction of species and the conversion of ecosystems—continues, as does human demand for land and resources as economies, human populations, and global wealth grow. In the lectures of countless conservation scientists, slides show graphs of number of species, population sizes, or the extent of habitat trending depressingly downward, sometimes with a counterprocess (say forest clearance, use of pesticides, or density of roads and settlement) rising inexorably, like a deadly mirror image.14 The catalog of destruction indeed makes dour reading. In terrestrial environments, grasslands, dry forests, and savannas are being converted for crops and degraded due to heavy grazing by domestic animals. Tropical forests are being felled for oil-palm plantations, soy farms, and cattle pasture. In the oceans, coral reefs are dying as seas warm and become more acid because of climate change. Plastics are showing up everywhere, including in the benthic zone, thousands of meters down. The Ellen MacArthur Foundation calculates that by 2050 plastic waste in the oceans could weigh more than all the fish.15 Large marine predators suffer from chemical pollution, and many large predatory fish, so sought after for human consumption, are severely depleted in number. Trade for bushmeat, pets, and wildlife products such as ivory has emptied many ecosystems of rare and large-bodied species. Across large swaths of country, the abundance of once-common birds is declining. In India, for example, of 867 bird species for which there are long-term data, over half had fallen in numbers since 2000, with 22 percent declining strongly.16 The form that terrestrial ecosystems take, and the dominant biological and physical processes that shape them and shape life within them, are now almost everywhere directed by human decisions about land use and ecosystem management. At the turn of the millennium, a new term was coined by the Nobel Prize–winning chemist Paul Crutzen to express the scale of these changes. He suggested that we live in a new geological era, the Anthropocene, in which humans are the dominant biological and physical force shaping the earth.17



The Place of Nature 9

The landscape around the ancient woodlands of eastern England is a classic product of the human propensity to take and remake nature to yield a good return. Although rural, it is highly industrialized. In spring, the small woodlands are filled with birdsong, places of shadow, and shafts of sunlight. The fields around them are uniformly bright and, except when crop sprayers are at work, completely quiet, empty of visible life beyond their growing crops. Few birds or pollinating insects move across the neat crop canopies, which stretch for half a kilometer without a break. This landscape, from which hedges and trees have mostly been removed, has been fashioned into what the UK farmers sometimes describe as their “factory floor”: a place designed and managed to maximize agricultural efficiency. It is beautiful in its productive way, with its even carpet of crops stretching to the horizon, each growing at the same speed, each reaching maturity at the same moment, each variation in soil fertility or crop health standardized by automated delivery of fertilizer and pesticide in exact proportions to optimize the growing crop, with GPS-linked tractors and detailed satellite and drone maps of the crop condition. It would be easy to conclude that the boundary between the nature reserve and the arable field marks the boundary between a world of nature and a world where the idea of naturalness has lost its meaning and is uncontested. But that would be highly misleading, for these silent and tightly organized fields have been the epicenter of a quite different kind of concern with naturalness. In July 1999, members of the environmental campaign group Greenpeace entered a six-acre field near the city of Norwich, just an hour’s drive from Cambridgeshire’s ancient woodlands. The field was growing an experimental crop of genetically modified (GM) maize (corn). Genetic modification (GM) of that era involved the insertion of genetic material from one species into another, often using yet a third (often a virus) as a vector. In the case of crops, this was usually done to improve yields or increase pesticide or herbicide resistance (allowing, for example, grass crops like maize to resist herbicides formulated to kill grass weeds, as in Monsanto’s “Roundup Ready” corn). This crop was part of a discreet government-sponsored trial to determine whether the GM seed could be sold to farmers for use in the United Kingdom.18 Arriving secretly in the early morning, Greenpeace protestors dressed

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The Place of Nature

in white hazard suits entered the field and used a mowing machine and strimmers (weed-eaters) to harvest the growing corn crop. Farmworkers tried to drive them away with a mechanical digger, and the farm manager disabled the mower by ramming it with a tractor. The police were called, and thirty people were arrested. So far, a classic protest. The agrochemical company carrying out the trial, Aventis, condemned the protest as a “deliberate act of trespass and criminal damage upon private property.” For its part, Greenpeace said that “the protest had been a peaceful action, on behalf of the British public, against . . . ‘genetically-­ modified pollution.’” Meanwhile the UK government said that it needed field- and farm-scale trials to ensure that decisions about GM crops were taken on the basis of the best scientific evidence.19 Twenty-eight of the Greenpeace activists were prosecuted, charged with stealing the corn crop. A trial in April 2000 acquitted them, accepting that their intention had been to harvest and bag the corn and return it to Aventis, which owned the crop. A second trial in September 2000 cleared the protestors of a second charge of causing criminal damage. The jury accepted that they had a lawful excuse for their actions, since under the Criminal Damage Act of 1971, property can be lawfully damaged if the action is to protect other property, and the jury accepted the Greenpeace contention that the maize was attacked so that its pollen could not “genetically pollute” nearby crops: the action was therefore legal.20 Greenpeace was triumphant, and the agrochemical company, farm owner, and government were cast low. In commenting on the case, the UK government and the company both criticized the protest from the perspective of science. The government said their top priority (ironically just like the protestors) was “to protect the environment and human health.” Without “strictly controlled research,” they argued, widespread GM crop planting would take place without “real scientific evidence” about GM crops. They announced that GM trials would continue. Meanwhile, the organization representing the agrochemical companies (the Supply Chain Initiative for Modified Agricultural Crops, SCIMAC) told journalists, “We’re disappointed that an extremist minority didn’t have enough confidence in the scientific strength of their own arguments to let the science decide.”21 This is an old story, just one bitter episode in the messy and (thus far)



The Place of Nature 11

mostly unsuccessful campaign to gain public acceptance for genetically modified crops in Europe. The European Union (EU) put a de facto moratorium on new approvals of genetically modified organisms (GMOs) in 1998. But meanwhile, many GM crops became standard in the United States and other countries, to the extent that three-quarters of processed food consumed in the United States contains one of eight GM crops (corn, soybean, alfalfa, sugar beet, canola, cotton, papaya, and squash). Furthermore, the European ban was soon undermined by global trade. Without knowing it, consumers began to buy and eat products from GM plants, because EU countries imported them in processed form (particularly soy for animal feed). An increasing number of exemptions were granted by individual national EU governments allowing GM seeds to be planted.22 The outcry over GM crops may not seem very relevant to the challenge of declining nature and its conservation. After all, if nature is safely tucked up in its reserves, how relevant is the idea of naturalness to the question of what happens outside them, where nature has been declared by many to be absent? However, the idea of naturalness proved central to the debate about the genetic novelty of the new GM crops. And the question of how novel GM crops are turns out to be central to the questions about wild nature that are the focus of this book. Leaving aside, for the moment, important issues such as the risk that modified genes might spread to crops’ wild relatives, or the way corporate inventions were tested and approved, what is really interesting about both this story and the court case is the way they drew attention to the genetic makeup of the corn crop and the way its genes had been manipulated. Controversy over this issue continues, in both the United States and Europe and many other countries around the world. For example, in 2015, GM seed was found in a batch of imported conventional canola (oilseed rape) being grown in experimental sites in England and Scotland as part of official registration of new plant varieties. The UK government’s GM Inspectorate announced that the seeds would be destroyed. Likewise, salmon-colored petunias planted throughout Europe and the United States turned out to be genetically modified with a gene from corn inserted to give the vivid coloration and were all pulled from the marketplace because regulators had never given permission for their sale.23 The question of the “purity” of seed offered to farmers and gardeners

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The Place of Nature

by the agricultural supply industry involves an argument about nature. Whatever one thinks of it, the public and official concerns about the genetic makeup of genetically modified plants turns on the question of whether they were in some way unnatural. This is odd, because the unnaturalness of the drastic ecological simplification and intense management of commercial agriculture had been acknowledged (and argued about) for decades. It was the unnaturalness of the shocking and unfocused killing power of organochlorine pesticides that gave Rachel Carson’s Silent Spring its cutting edge. The removal of not only pests but also nontarget species from farmland by commercial agriculture’s chemical regime contributed to the need for nature to be protected in reserves. So why worry about what is “natural” in what is by any standards a very unnatural system—the modern agricultural field?24 Opposition to GM crops turned not on the “unnaturalness” of an ecosystem drenched in pesticides and artificial fertilizer, because almost all fields were like that. The campaign focused strongly on the question of whether the GM crops themselves were “natural.” Campaigners coined the devastatingly effective term “Frankenstein foods” (or “Frankenfoods”) to persuade the general public that in some way they were not. Despite the lack of conclusive evidence of direct human health impacts of consuming foods made from GM plants, this term spread among environmentalists, food activists, and worried consumers, in the United States, the United Kingdom, and more widely. Use of the term “Frankenfoods,” first deployed in the United States in 1992 in campaigns against the Flavr Savr tomato created by the California company Calgene, has subsequently spiked at different times in different countries, as debate about GM technologies has waxed and waned.25 The co-option of Mary Shelley’s Frankenstein in this debate is significant. In the story, Dr. Frankenstein is, of course, the maker of the animated humanoid—the creator, not the creature. The creature remains unnamed, and except for his aesthetic challenges, his uncanny speed and endurance, and (eventually) his vengeful spirit, he is in some ways a more humane being than his maker. Indeed, in many ways the accusation of monstrous character is better directed at the creator and his experiments and not the creation—despite common readings of the plot of the novel and films inspired by it.



The Place of Nature 13

The allusion to Frankenstein’s creature in the context of GM crops was designed to make the work of crop geneticists and seed companies seem unnatural. It was effective at one level, in that the creature was certainly artificial, being assembled from cadavers in Frankenstein’s laboratory and animated by galvanic shock (creating a classic scenario for the gothic-­ novel genre that grew from Mary Shelley’s work). It is in the artificiality of Frankenstein’s creature, the unnaturalness of his creation and of the mind that conceived it, that the story’s horror has its root. The question of naturalness cannot be contained within a simple landscape-­ scale juxtaposition of nature reserve and field and the modified corn planted there, whether on the level of a European agricultural landscape, a rain forest, or a city. The allusion to Frankenstein shows us that concerns about naturalness extend from the ecosystem down to the level of the body and its composite elements. At that level, within the corn plant and the human body, people have inadequate frames within which to think about what naturalness means. In desperation, they turn back to Frankenstein the experimenter and his unfortunate and unhappy “monster.” Humankind needs a better way to think about naturalness in the context of nature and conservation of diversity. Industrial field and protected nature offer a crude binary distinction, a territorial divide, separating the humanized and natural domains. This division has enormous significance, both for the way the environment is managed and for the way nature is imagined. Protected areas may be tiny and human-managed, like Hayley Wood, or vast and relatively undisturbed, like the Pantanal in Brazil, or Greenland. But whatever their size, in our minds there lies nature: beautiful, diverse, valuable, and free from human influence. Outside them we imagine the spreading human world: developed, artificial, and unnatural. The juxtaposition of nature reserve and field seems to mark a fault line in our ideas of how to live with nature. In conservation, thinking about nature has been dominated by the concepts of ecosystem and the species. In proposing the ecosystem concept in 1935, the British ecologist Arthur Tansley suggested that “the organisms and the inorganic factors alike are components which are in relatively stable dynamic equilibrium.” Half a century of popular environmentalism has given the concept of the ecosystem universal currency. The categories

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The Place of Nature

of coral reefs, prairies, rain forests, or reed beds are familiar, as are accounts of ecosystem loss and conservation and the language of “ecosystem services,” such as clean water or flood protection. The existence of variety within each ecosystem category is intuitively recognized: rain forests with orangutans in Borneo are not the same as rain forests with howler monkeys in the Amazon or rain forests with gorillas in the Congo Basin of Africa.26 The species has always been at the heart of conservation. The founders of late Victorian conservation organizations like the Society for the Preservation of the Wild Fauna of the Empire (now Fauna and Flora International) were motivated by stories of the extinction of animals like the passenger pigeon (still present in their billions in North America earlier in the nineteenth century), or the quagga (a form of the common plains zebra, exterminated because it ate pastures reserved for white farmers’ sheep in the South African Cape), and the rescue of others (such as the American bison).27 Species remain the best-studied dimension of biodiversity and at the heart of the conservation concern. The category has a long-established scientific heritage in 250 years of taxonomy and is simple and widely recognized. Injunctions to “save the whales” or “save the rhino” are familiar themes in the language of conservation. Public appeals for donations often build on the need to save endangered creatures, showing a predilection for larger animals (pandas, elephants, or gorillas). Although conservation organizations recognize the dangers of bias toward charismatic species to the exclusion of others, there still tends to be a disproportionate focus on mammals, birds, amphibians, butterflies, and flowering plants, rather than insects, mosses, or fungi, let alone microorganisms, which form an integral and vital part of living diversity.28 As with ecosystems, variety within each species is important in conservation. For example, different races of gray wolves across the world look distinct from one another, not only in color but other characteristics (for example, Mexican wolves are slighter and shorter haired, while Mongolian wolves are larger and have shaggy coats). These differences are often honored by subspecies names and incorporated into many species conservation programs. Species and subspecies are both accepted as natural units for conservation efforts. Thus, the Amur leopard of the Russian Far



The Place of Nature 15

East, although only a subspecies of the very broadly distributed leopard (which ranges all the way to southern Africa), is distinctive enough to have its own conservation program. One critical element of biodiversity is often overlooked. Most organisms existing on earth are composed of only a single cell: microbes, a category that includes bacteria, archaea, fungi, algae, protozoa, and viruses. Humans are visual animals and microbes are small. As Ed Yong writes, “Bacteria are everywhere, but as far as our eyes are concerned, they might as well be nowhere.” Yet microbial diversity is incredible. In the backcountry of Yellowstone National Park (noted for its wolves, bison, and elk), in an area rich with volcanic activity and hot springs, Daniel Colman of Montana State University and colleagues found a community of microbes they described as “incredible, unique and truly weird.” Whereas most hot springs contain only a few types of microbial organisms, this spring, named Smokejumper 3, had representatives from almost half of all known groups of microorganisms, including many that had not previously been described. There was more diversity in one small sample of water from Smokejumper 3 than all of the animal and plant biodiversity found in the rest of Yellowstone.29 Diverse communities of microorganisms are found floating in the air high above the earth, tossed by ocean waves, in the soil, on the skin, and in the bodies of all plants and animals. Microbes have even been recorded 750 meters below the seafloor in the Indian Ocean. The richness and diversity of microbial communities make them important conservation targets in their own right. Microbes are vital to the functioning of all parts of the earth’s living systems—essential to the ocean’s carbon cycle and to the nitrogen cycle in grassland soils. They comprise part of the coral polyps making up the world’s great tropical reefs. They are also the constituent parts of the microbiomes that lie within and on the bodies of many of their larger and more structurally complex cousins—including humans, frogs, and plants.30 Not only are there a lot of different species in a microbiome, they interact with host cells in a remarkable number of ways. In humans they are thought to influence health problems like obesity, circadian rhythms, diabetes, and perhaps even some forms of cancer. The microbiome of a cow’s intestine consists of an astonishing seven thousand to ten thousand

16

The Place of Nature

species of bacteria, archaea, protozoa, fungi, and viruses, interacting with the cow’s physiology, influencing digestion and immunology. Many plants have microbiomes on their roots, on the surface of the aboveground plant, and within the plant tissue. These influence germination, growth, nutrient uptake, disease resistance, and stress tolerance and can affect dispersal, competition, community structure, and ecosystem function.31 However, the most fundamental element in nature’s diversity exists at a smaller scale still than microbes. It is the gene—the coils of DNA that give shape and structure and life to all living organisms. The gene, or the collective genome of an organism, is the too-often-ignored actor of the conservation world. It does its work unseen, unremarked, and underappreciated. Genes are essential to the way organisms function. They help shape the species and ecosystems that they make. Without genes, biology simply doesn’t work. It is not that the biological importance of genes is unknown—in most university biology departments, molecular biology massively outweighs organismal and ecosystem biology. Moreover, conservation genetics is a well-­established field, although it is small compared to conservation biology or ecology. Relatively few conservation scientists and practitioners are well versed in the genetic dimensions of their work, and the science of conservation genetics barely features in the work of large conservation organizations. Relatively few biologists focus their work on the genetics of wild species and the ecosystems within which they exist.32 Conservationists have long recognized that the terms “species” and “ecosystems” are just convenient shorthand for the diversity of the natural world. When the Convention on Biological Diversity fixed the definition of biodiversity in 1992 as “the variability among living organisms,” it deliberately included diversity of ecosystems, diversity between species, and diversity within species. The diversity of the myriad of ecosystems, species, and genes and their interactions make up the world’s biological richness and is responsible not only for the functioning of nature as it now exists, but also for its ability to adapt and evolve under changing conditions. The 1992 convention required signatory governments to take steps to preserve all the variety of life, ecosystem, species, and genes. But even though the definition of biodiversity explicitly incorporates



The Place of Nature 17

the genetic, diversity at the level of the genome remains little thought about. After all, you can’t grab your binoculars and go gene watching, or imagine getting lost in a three-day trek through the genome. The variety of life at the genetic level is, for many people who care about nature, out of sight and out of mind. It has virtually no public constituency. There are no citizen groups lobbying to save genes—yet. If the future of the genome of wild species is still to an extent terra incognita for most people in conservation, wider questions about genomes are being asked, notably in public concern about genetically modified crops. There is something of a mismatch between the level of interest in genetic dimensions of human medicine and in GM food, and in the relative lack of attention given to the genetic dimensions of nature and its conservation. Scientific discoveries are bringing these issues together. The new field of “synthetic biology,” the “design and construction of novel artificial biological pathways, organisms and devices or the redesign of existing natural biological systems,” is ushering in a new world, fraught with potentialities and pitfalls. Synthetic biology allows rapid changes to the genetic makeup of animals, plants, and other organisms. Genomic changes that evolution once brought about over many millions of years can now be made in a laboratory in months or days.33 Such work has profound implications for wild species and the practice of conservation. New techniques, described as “gene editing,” allow specific and detailed changes to be made in the genetic code that shapes the bodies of organisms, from simple microbes to the complex mammals. These techniques are more sophisticated and more powerful than the old techniques of genetic modification. They give scientists an unprecedented ability to design and make precise genetic changes to living organisms. Such changes are already being made to domesticated animals and crop plants. Work is also beginning to be done to alter the genomes of wild organisms. This is already happening as part of agricultural pest control. In 2017 a team of scientists led by Anthony Shelton of Cornell University released a genetically engineered moth into a cabbage field in New York State. The species was the diamondback moth, an unassuming small brown insect. This moth does not fly particularly well, and yet it is one of the

18

The Place of Nature

most widely distributed moths on the planet, moving on wind currents. Like many other world-bestriding species, it has also piggybacked the spread of people, and its caterpillars are a serious pest on crops in the mustard family, such as canola, cabbage, broccoli, kale, and Brussels sprouts. The damage it causes is said to amount to $4 billion or $5 billion a year.34 As a key economic pest, the diamondback moth has been the target of pesticides for many decades and has acquired resistance to many of them. So scientists have begun to experiment with other ways to attack it, by altering its genome. Working with a British biotechnology company called Oxitec (originally a spin-off from the University of Oxford), the Cornell team genetically engineered a strain of diamondback moths so that they produced only male moths. When these were released, wild-type females that mated with them were not able to produce viable female offspring. Greenhouse experiments showed that the moth population was suppressed, and the insecticide-resistant genes were diluted. When released into the wild, in a field on Cornell University’s New York State Agricultural Experiment Station, the engineered male moths survived between three and five days, and moved about thirty-five meters from their point of release. The potential of genetically engineered “self-limiting” moths seemed proven.35 This experiment, conducted under a permit from the US Department of Agriculture, was the first open-field release of a genetically engineered animal in the world. It is just one of a very large number of applications of new technologies to reengineer genes and genomes. Many applications of these technologies focus on biotechnology applications, envisaging organisms with modified genomes that will permanently be enclosed in vats or industrial facilities. Increasingly, there is talk of releasing them directly into the open environment, like the diamondback moth was. Conservation applications of gene-editing techniques of this sort are also being considered. One recent experiment, for example, addressed the sharp decline in the honeybee, a key provider of pollination services in many parts of the world. The declines are thought to be partly the result of the interaction between the parasitic varroa mite and a virus with the ugly name of “deformed-wing virus.” Researchers at the University of Texas at Austin selected a bacterium (Snodgrassella alvi) from one of the nine species clusters in the microbiome of the honeybee’s gut and engineered its DNA. This caused the release of double-stranded RNA mole-



The Place of Nature 19

cules that repressed host gene expression and altered the bee’s physiology, behavior, and growth. They showed that this improved bee survival after being challenged by the virus, and that the engineered bacterium could kill parasitic varroa mites by triggering their immune response. A commentator described the work as “A microbiome silver bullet for honeybees.”36 The revolutionary potential of the field of synthetic biology, which only began in the new millennium, has had an astonishing impact on the imagination and ambition of scientists, drawn to it both from computer engineering and biology. The American synthetic biologist and computer engineer Tom Knight expressed the scientists’ excitement, describing genes in the language of software, saying, “the code is 3.6 billion years old; it’s time for a rewrite.”37 True to such vision, genetic editing is being applied today in a wide range of fields, in laboratory science (particularly human medicine), in agriculture (crop breeding and pest research). and in industrial biotechnology (food production, fragrances, and pharmaceuticals). Engineering of the genome is powering a new economy of life, and the rate of innovation, investment, and growth is prodigious. In all these sectors, and at the core frontiers of biological research itself, science has found a way to unlock the genome and recode it to perform in new ways. Conservation scientists are also starting to explore the potential of these new techniques to contribute to their understanding and their work. At the same time, they do have a particular awareness of the potential risks of this kind of experimentation. Conservation needs to decide how to respond to synthetic biology. The ability to edit genomes muddles nature and artifice in new and profoundly challenging ways. Faced with its powers, and the challenges, opportunities, and risks it brings, the critical question is this: How should we think about nature in a world of genome editing? That is the question we answer in this book. First, we have to ask what nature is, and what naturalness means.

2 The Problem of Nature

At dawn a straggle of young whooping cranes fly in a ragged “V” behind an ultralight aircraft. It is their annual migration, and they are being taught the route by a human pilot dressed head to toe in white. This bizarre sight is one element in a complex conservation program created to try to bring the whooping crane back from the edge of extinction.1 Whooping cranes are extraordinary birds. They stand up to one and a half meters (nearly five feet) tall. Their plumage is a gleaming white, with a striking smear of black on the face and a red cap on top of the head. They breed in wetlands up in the muskeg and taiga of Canada and the northern United States, and every year they fly south to winter along the Gulf of Mexico. But by the middle of the twentieth century, they were almost extinct. Whooping cranes were never numerous, with perhaps ten thousand birds before Europeans reached North America. Hunting drove them close to extinction. There were still perhaps fourteen hundred birds in 1870. But by the end of the 1930s, there were just fifteen adults in a single migratory flock, plus a few more that had taken up residence in Louisiana. Intensive conservation efforts began, centered on captive breeding. By 1976, the wild population was still only sixty birds, all breeding in Wood Buffalo National Park in Canada. Conservation efforts were redoubled.

Whooping crane 20



The Problem of Nature 21

Captive cranes did not mate readily and proved poor parents, so staff forced artificial insemination and removed eggs from females (to encourage multiple egg laying), using other species or machines to incubate them. The most extraordinary aspect of this protracted effort (and financial investment) in species recovery was the invention of the “human crane.” Once hatched, crane chicks were taught to eat and drink by people hooded to conceal their shape and carrying arm puppets designed to look like adult cranes. But the genius was in what came next.2 In the wild, cranes learn migration routes by following adult birds, with whom they stay for eleven months. The whooping crane chicks were trained to follow an ultralight and taught a migratory route across the North American continent. This approach was well established with geese, which had readily imprinted on human caretakers. Experiments with cranes began in 1995 in Wisconsin. Ground-based training with ultralights was followed by several months of training flights. Finally, the aircraft-led southward migration began, with the young cranes guided in stages over nearly two thousand kilometers from Wisconsin in the northern United States to a wintering ground in Florida.3 Having taken the trip once, the cranes flew back north in spring on their own, and then repeated the migratory path the next year and the next. Radio-tracking showed that over the next few years aircraft-trained birds changed their migration patterns, copying birds that had learned their routes from other birds: migration is a culturally learned behavior, and hand-reared cranes go on learning. But the technique worked. By 2017, 505 whooping cranes migrated to Arkansas National Wildlife Refuge, including 49 juveniles, and with captive birds, the world population had risen to over 800 birds.4 Thus far, this reads like a classic conservation story, a species saved from extinction through the diligent application of time, money, and hard work. Change was to come. In 2017, the whooping crane captive breeding program, at the Patuxent Wildlife Research Center in Maryland, closed after fifty-one years, when government funding for the $1.5 million program was cut. The writing had been on the wall for a while. In October 2015, the US Fish and Wildlife Service published their strategic vision for the eastern migratory population of the whooping crane, proposing an end to ultralight-guided migration and all costume-rearing methods. This

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The Problem of Nature

wasn’t because the conservation job was “done” (the crane population was still way below its historic level, and there were multiple threats to its survival on breeding grounds and on migration), or just that the federal government expenditure was being cut (although this was certainly an issue in 2017). The Fish and Wildlife Service had other concerns. It had decided that the use of ultralights was too “artificial.” It preferred methods that could “reduce artificiality and better mimic the natural conditions experienced by wild-hatched whooping cranes as closely as possible.”5 The language used tells us something about the way conservation decisions are often made. If we leave aside the issue of “value for money” (and the everlasting politics around the budget of US federal agencies), the idea that leaps out from this statement is that the real problem was “artificiality” of the way cranes were raised and trained to fly. This raises two obvious questions, neither of them easy to answer. First, What does it mean to say that a conservation method is “artificial,” or the obverse of this, that it is “natural,” and how do we distinguish between them? An ultralight aircraft is clearly a twentieth-century human invention, but airborne human guides had managed to teach hand-reared cranes their “natural” migration route. Does the migration of cranes on a route they were first taught by a human pilot always remain “artificial,” even after the aircraft is no longer being used? Does the adaptation of destination as a result of learning from other birds start to make the behavior more “natural,” and if so, what determines how fast artificiality bleeds away and naturalness grows in its place? The second question is equally tricky and more universal. It is this: How much artificiality is okay for conservationists to use to protect nature? As we have seen, the contrast between the naturalness of a woodland and the artificiality of gene-edited crops seems an obvious one. If there are questions about the degree of artificiality of using hand puppets and ultralights to raise crane chicks, how should we think about the reengineering of the genetic makeup of wild species? Should the methods conservationists use have to fit some standard of “naturalness”? We certainly expect our nature reserves to look natural. For example, buildings in US national parks specialize in a frontier-timber aesthetic, and paths are landscaped to be discreet in the landscape. Do we also expect the methods used by conservation to be natural? And if so,



The Problem of Nature 23

what do we mean by this? How much artificiality is acceptable, and how is it to be measured? These are not small questions. In describing the life on earth, it is very common to make a distinction between what is natural and what is artificial. Something (anything—a sea snake, a mushroom, or a mountain) is commonly described as “natural” because it is not the result of human work or ingenuity. By contrast, those things that bear the unmistakable mark of human manufacture (a smartphone, a waterwheel, or a city) are described as “artificial.” It is normal shorthand for conservationists to say that the construction of a mine, dam, or highway in a tract of tropical forest is a threat to a “natural” ecosystem. This simple distinction between natural and artificial is linked to a moral judgment. It is commonly held (very strongly so by naturalists and conservationists) that natural things are to be valued more than artificial things. People often say they prefer “natural” foods and distrust “artificial ingredients,” even if they don’t have a very precise idea of how their food is produced (as we have seen, public opposition in Europe to GM food turned on the sense of its “unnaturalness”). And settings such as nature reserves or woodlands are widely thought to have benefits for human mental well-­being, while built-up urban spaces do not. Even pictures of nature (trees, for example) are thought to benefit the recovery of hospital patients.6 The idea of nature as something separate from humanity, so pervasive in Western cultures, goes far beyond the thinking of conservationists. It has ancient roots. In describing the development of mechanical devices in ancient Greece in her book Gods and Robots, Adrienne Mayor shows how distinctions were drawn between what was “made” and what was “born.” She argues that the difference between biological birth and manufactured origin has marked “the border between human and nonhuman, natural and unnatural” ever since classical times.7 This distinction became part of the radical (and world-changing) agenda of rationality, empiricism, free thought, and progress that comprised the European Enlightenment in the seventeenth and eighteenth centuries. The observation of nature was central to the revolution in thought that pitched science and reason against the medieval idea of universal truth and religious authority.8

24

The Problem of Nature

To the amateur scientists of Europe’s new learned academies, nature was understood as something that could be rationally observed and measured and which, using that knowledge, could be managed and exploited in new ways. From the seventeenth century, new knowledge catalyzed intellectual inquiry and stimulated ambition to transform and “improve” nature and society across the vast terrains of new European empires. The logical separation of nature and society was fundamental in both unleashing the carbon-guzzling machines of the Industrial Revolution and imperialism’s drastic world-changing ambition. It underpinned the idea that nature was there as a resource, needing only rational exploitation to maximize human benefit. In time, it also allowed the understanding to develop that exploitation could damage the earth. From visionary works like George Perkins Marsh’s Man and Nature in 1864 grew the utilitarian conservation movement of the late nineteenth century (and, by circuitous routes, the sustainable development movement of the late twentieth). The idea of a nature separate from and threatened by humankind lay beneath the emergence of the modern conservation movement.9 Romanticism, rising in Europe in the late eighteenth century, was a reaction against both the scientific rationalization of the Enlightenment and the brute exploitation and degradation of nature in mines, factories, and slums. Nature began to be seen as sublime and pure, a source of renewal for the spirit and aesthetic experiences. In art, poetry, music, and literature, rural landscapes were presented as pastoral idylls, and wild scenes such as mountains thrillingly fearsome—literally “awesome.” The British philosopher Kate Soper argued that “untamed nature began to figure as a positive and redemptive power only at the point where human mastery over its forces is extensive enough to be experienced itself as a source of danger and alienation.” In Europe, wilderness had long been feared as uncivilized land, a world where wild beasts and lawless people dwelt. Romanticism and industrialization changed that.10 In the United States, in the nineteenth century, the idea of wilderness offered an antidote to industrial modernity. It struck a deep chord in an immigrant society, as the struggle to occupy the apparently endless Western frontier began to run out of space. Writers like John Muir created a whole new world of nature writing, celebrating the profound beauty and serenity of landscapes like Yosemite. As the distinguished environmental



The Problem of Nature 25

historian William Cronon pointed out in his influential essay “The Trouble with Wilderness,” the idea of wilderness is a profoundly human creation, specific to particular cultures and historical eras. It is a mirror, and as we gaze into it “we too easily imagine that what we behold is Nature when in fact we see the reflection of our own unexamined longings and desires.”11 These ideas now have global resonance. Everything from travel brochures to wildlife documentaries present nature as an “other” to which we can travel, physically on holiday or, indeed, virtually through media imagery. Wildlife documentaries often adopt a simple binary shorthand, equating the naturalness of an ecosystem with an absence of people and visible human handiwork. Wilderness always has to be imagined before it can be protected.12 In Europe, nature was valued slightly differently in the nineteenth century, but the influence of culture on ideas of nature were, if anything, more profound. In the United Kingdom, the beauty of nature was a key factor in the rise of the conservation movement, and the idea of natural beauty was applied unselfconsciously to land and landscapes that not only bore the unmistakable signature of human enterprise (such as the Lake District’s stone walls or grazed hills) but on a countryside whose character was chiefly formed by centuries of human management. In British conservation, the scientific importance of ecosystems and species is wrapped up in broader cultural values attached to some of those species (the nightingale, for example, hymned in a famous poem by John Keats) or spaces (for example, the childhood romanticism of the Wild Wood in The Wind in the Willows).13 Is it only in places like Europe, with a long-recorded history and published literature, that such cultural values are important? Not so, for ecosystems have been shaped by human occupation across the world, including the vast rain forests and drylands of South America, Africa, and Asia. With the exception of Antarctica (which humans never settled, even though legions of tourists now visit every year), regions that seem wholly natural have almost all been lived in by somebody. We may have no formal written record of the cultural values these people attached to the ecosystems in which they lived and died, but it is unreasonable to assume those values did not exist just because no trained scholar wrote them down. The world’s great “wildernesses” are cultural spaces as much as natural.14

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The Problem of Nature

The complexity of the idea of nature has long been recognized by philosophers. As the novelist and literary critic Raymond Williams pointed out in his book Problems in Materialism and Culture, people write about “nature” as if its meaning were given, but in fact the word carries a wide range of meanings. Nature may be said to be violent (“red in tooth and claw”), reflecting the idea of a ruthless competitive struggle for existence. But it can also be thought of as self-regulating, an interlocking of mutual advantage and cooperation. Ideas about nature are always, as Williams put it, “the projected ideas of men.”15 Today, the word “natural” is used in several ways. First, the word is conventionally used to describe something that lies beyond human artifice— something subject to its own rhythms and dynamics. This kind of naturalness exists independently of the observer. It can be measured and analyzed by natural scientists, whether in a physics laboratory or on the Greenland ice sheet.16 Second, the word “natural” can also be used to refer to an essence, something intrinsic and unchanging (a fox, for example, said to be “wily,” or a breakfast cereal described as being full of “natural goodness”). Human behavior is often described as “natural”: it is deemed natural for parents to love their child, or natural to grieve a lost relative. In the same way, behavior (of people or animals) can be described as unnatural, for example, a fox eating out of a trash can, or a human showing an “animal-like” appetite. Finally, we also describe elemental forces as natural. Volcanic eruptions, typhoons, or floods are referred to as “extreme natural events.” We describe dynamic ecosystems that are unmodified and unconstrained by human actions as “natural”: natural rivers, for example, are understood to be those without flood-control dams; natural coasts lack seawalls; and natural fire regimes are those in which fire burns independent of human agency. All of these uses of “natural” reflect cultural determinations. They reflect how people in a particular culture at a particular time choose to speak about each other or about the living and nonliving world around them, rather than (necessarily) the attributes those things actually have. So an ecosystem might be described as “natural” without either speaker or listener having specific historical knowledge of the form and impact of human influence upon it. It makes perfect sense in normal life to say that the park



The Problem of Nature 27

is more natural than the town square, or that a river is made more natural by removing its concrete banks and letting rushes grow. However, this is more problematic when the same language is used for more technical purposes, for example, when discussing the conservation of ecosystems. How natural is it to teach cranes to fly with an airplane? Is there a limit to how artificial conservation methods can become before some critical element of naturalness is lost? That question is a fault line that runs right through contemporary thinking about conservation and is one of the structuring issues of this book. The instinct to preserve dominated the early conservation movement of the late nineteenth and early twentieth centuries, growing out of the collection mania of aristocrats, national museums, botanic gardens, and zoos. It remained important through the twentieth century in the establishment of national parks and other protected areas, designed to protect a representative sample of ecosystems and biomes.17 The idea of nature in terms of a series of “objects” (shrinking ecosystems and threatened species) was useful. Historically, natural environmental changes seemed relatively slow compared to rapid and destructive human-driven change. It continued to provide a basis for success as conservation planning became increasingly scientific and systematic in the 1990s, with satellite imagery and computers allowing the development of prioritization algorithms, and the definition of hotspots, important bird areas, and ecoregions.18 The trouble with this approach is, as ecologists have long known, that nature is not static but is subject to constant change. Conservationists face a moving target, a nature that shape-shifts and changes. Onto this they somehow have to fix the classifications and categories that allow them to identify problems and make plans to address them. Once, the tension between the understanding of nature as a set of static objects and nature as a moving kaleidoscope of species was less acute. In his 1905 book, Research Methods in Ecology, the American ecologist Frederick Clements argued from his fieldwork on the shores of Lake Michigan that vegetation communities developed through time to a stable endpoint, a “climatic climax” that was characteristic of each location. Arthur Tansley’s concept of the ecosystem in 1935 was more dynamic, embracing

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The Problem of Nature

soils, physical processes, and human action in “relatively stable equilibrium,” although it still allowed classification into characteristic “types.”19 The rise of theoretical and mathematical ecology, ecological energetics, and systems ecology after the Second World War turned attention to the quantitative interactions and dynamics within ecosystems. But even so, ecologists still tended to work within an assumption that ecosystems tended toward an equilibrium state, reflecting the idea that there was an inherent “balance of nature.” Ecologists used the language of machines, of feedback and control, to describe the way ecosystems self-regulated, arguing that if they were disturbed, they would tend to return toward an equilibrium state.20 This “classical” or “equilibrium” paradigm in ecology dominated ecology (and conservation) until the 1970s, when ecological modeling began to reveal complex dynamics and multiple stable states. Ideas about resilience and disturbance ecology emphasized the importance of patterns of disturbance and the irregular dynamics of ecological processes, including shifts between alternating stable states (for example, in lakes). There was renewed emphasis on the ways in which the physical and biological processes that cause ecosystem change vary in space and time and, hence, on the importance of context, contingency, and scale in the evolution of ecosystems: the ecosystem offers opportunities and constraints within which organisms live, but those organisms also shape and regenerate that environment.21 Changes in the science of ecology have profound implications for conservation management. Ecologist Daniel Botkin asked rhetorically in his book Discordant Harmonies, “How do you manage something that is always changing?” Past disturbance events (for example, tree falls, disease epidemics, or floods) are of fundamental importance to the present composition of ecosystems and, hence, to the “natural” features of conservation concern. The frequency, intensity, and scale of disturbance events are determined by the interaction of multiple factors, including climate, weather, fire, topography, geology, and the almost infinite interactions among the species present. Increasingly often, disturbances also reflect human agency, making the history of management critical to understanding contemporary ecological patterns.22 In nonequilibrium ecology, nature is understood to be dynamic and



The Problem of Nature 29

highly variable and its trajectories through time to be open-ended and changing, not always tending to return to some past equilibrium point. Natural disturbance is part of the “nature” with which conservation is concerned, and human actions are an integral element in ecological change.23 Gone, therefore, are the days when conservationists could conceive of “nature” in equilibrium and portray human-induced changes in those ecosystems as somehow “unnatural.” Gone, too, are comfortable certainties about naturalness and the management regime needed to sustain it. Conservation scientists know their ecology and recognize that ecosystems are dynamic. And yet, as ecologists Buzz Holling and Gary Meffe pointed out in 1996, a great deal of conservation work is predisposed to methods of “command and control.”24 Throughout human history, societies have controlled and exploited nature for profit or human benefit (sometimes both together), through agriculture, forestry and fishing, hunting and gathering, and disease control. Ecosystems have been rationalized, simplified and refashioned to suit human ends. Conservationists usually see themselves as trying to push back against these processes. Yet, surprisingly, conservationists take much the same approach to the conservation of nature as do those exploiting it. They routinely cut, burn, poison, shoot, and trap wild species in the name of nature protection, seeking to control the forms nature takes, making direct interventions to shape populations and ecosystems, to try to increase the numbers of a threatened species or eradicate an invasive species, to limit fire here while encouraging it somewhere else. Even areas designated as “wilderness” are in receipt of constant management. In the United States, the 1964 Wilderness Act defines wilderness as “an area where the earth and its community of life are untrammeled by man, where man himself is a visitor who does not remain.” Despite this pronouncement, a great deal of management is done: a survey in 2016 found that management interventions had taken place on 37 percent of wilderness areas between 2011 and 2015. The most common management was of vegetation (planting, cutting, applying herbicide) and wildfire (fuel management, suppression, control lines, application of fire retardant) or postfire restoration, such as planting or applying soil and mulch. Less than a quarter of proposals for intervention were rejected.

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This is not a phenomenon confined to the United States: the vast protected areas of Australia are given constant management to counter key threats to endangered species, such as fire and invasive species.25 A lot of conservation effort goes into attempts to stop ecological succession, burning or cutting vegetation and sometimes applying herbicides. Vegetation succession has been recognized as a perfectly natural process since the early years of plant ecology at the start of the twentieth century (classically, open fields turning to scrub and to woodland, or open water developing fen ecosystems and eventually wet woodland). The trouble is that some species and sets of species only thrive in very specific conditions that only occur at particular points on a successional cycle. Not every rare species thrives in a mature forest, and to preserve some of them, conservationists have increasingly borrowed the methods of farmers, ruthlessly cutting scrub, and of foresters, felling timber to allow smaller trees to grow. A good example is the battle to conserve the Kirtland’s warbler, also called the jack pine warbler, in the United States. This small bird breeds in the pine forests of Michigan and migrates every fall to winter quarters down in the Caribbean. Once common, it is now rare. The problem is that it is very picky about where it will nest: only on the ground in dense thickets of young jack pine. Like many other pine species, jack pines are ecologically adapted to periodic fires. Indeed, their cones only open to release their seeds when the forest burns. Once, fire would rip through the forest every few years. The next year the new seeds would sprout and forest succession would restart again. Cue a nesting opportunity for the Kirtland’s warbler, which would move in about six years after a fire, when new tree growth was dense. After about fifteen years, the trees would have grown too big and the Kirtland’s warbler would abandon the burned area and move to another. Once jack pine forests were extensive and wildfire was a natural disturbance process that created a mosaic of mature and successional stages. Kirtland warblers could simply move about to find the new growth they needed. No more. Today there is little jack pine forest, and for decades nobody was willing to see it burn in wildfires. As a result, the few pine forests that remained underwent ecological succession to mature forest. In the breeding season of 1951, a census found only 432 singing male



The Problem of Nature 31

Kirtland’s warblers in the United States. Conservationists swung into action. In 1957 the Michigan Department of Natural Resources set aside three areas as warbler management units, and in the 1960s the US Forest Service wrote a species management plan and set aside sixteen hundred hectares of forest as a reserve. In 1973, the warbler was listed under the Endangered Species Act. Efforts were also started to trap brown-headed cowbirds (which lay their eggs in the nests of other birds) and remove them from warbler nesting areas.26 None of it worked. A census in 1974 recorded only 167 singing male warblers. Finally, research revealed the Kirtland’s warbler’s very specific habitat requirements, and a recovery plan was put in place to create them artificially. This involved clear-cutting mature jack pine and manually or mechanically planting pine seedlings to create a mix of thickets and scattered openings. Conservation managers left dead standing trees and small strips and islands of live trees during clear-cutting, to mimic conditions after a wildfire. Today, seventy-seven thousand hectares of jack pine are managed on a forty-year rotation, with about fifteen hundred hectares replanted every year. This creates about fifteen thousand hectares of habitat suitable for warbler nesting each year. Gradually, prescribed fire is also being introduced into habitat management. And this, at last, is working. Over 1,000 pairs of Kirtland’s warblers were recorded in 2001, and in 2007 they were reported breeding in Wisconsin and in Ontario, Canada. In 2018, the US Fish and Wildlife Service proposed removing the Kirtland’s warbler from the list of endangered species.27 So a success. But to save the Kirtland’s warbler, the forest had to be put to the torch and chainsaw to reestablish the dynamics of their nesting habitat. The Kirtland’s warbler is a classic example of what J. Michael Scott and his colleagues have called a “conservation-reliant” species—that is, a species that needs permanent life management intervention (for example, of habitat, predators, or parasites) to survive. They calculated that 84 percent of the US endangered and threatened species with recovery plans were conservation reliant. Without intensive artificial human management, many species would disappear. The harsh roar of the chainsaw has become the sound of nature being preserved.28 The desire to control natural processes is not confined to vegetation. Many conservation projects place great importance on the control of pred-

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The Problem of Nature

ators, installing extensive fencing, employing shooters to kill foxes or other predators with night sights, humanely trapping feral species such as mink and humanely killing them. Poison is also an important conservation tool. On the remote subantarctic island of South Georgia, for example, introduced brown rats and house mice were the dominant predators of the island’s seabirds and the ground-nesting endemic South Georgia pipit. In 2011, the South Georgia Heritage Trust began a campaign to spread poisonous bait on an industrial scale, with helicopters scattering poison on every meter of vegetated land. The exercise cost about $11 million at 2015 prices ($104 per hectare). By July 2017, twenty-eight months after baiting had finished, there was no sign of surviving rodents, other than one newly introduced, apparently, by ship in October 2014.29 Such methods have become a standard part of conservation tool kits, and not just on oceanic islands beset by unwelcome exotic rodents. In New Zealand, for example, the common brushtail possum (introduced from Australia in the 1850s as a source of fur) causes catastrophic ecological damage by eating the eggs of endemic birds, invertebrates, and plants. Conventional methods of possum control work to a degree but are either controversial (poison is unpopular because of fears about water supplies and the accidental poisoning of domestic dogs and introduced deer, valued by hunters) or expensive (traps are time-consuming to set and monitor in remote areas). In response, something of a growth industry developed making automated machines to kill possums and other predators. One company markets a self-resetting possum trap that can kill twelve possums one after the other by firing a piston into the back of the animal’s head when it bites a bait block. Another advertises a trap that can operate unsupervised for up to one hundred cycles or twelve months without servicing and bait refill. With the same intentions, a device has been developed in Australia to automate the killing of feral domestic cats by coating them with toxic goo that kills them when they groom themselves.30 The lengths to which conservation managers will go to exercise the control they believe to be needed is well shown by the management of African elephants. Bill was involved for some years in a project in Laikipia District in Kenya, which looked at conflicts between free-living elephants



The Problem of Nature 33

and smallholder farmers. Wild elephants have declined catastrophically in Africa as a result of poaching. Yet, where they survive, they have less and less land to live on, as agriculture has expanded and intensified on the lands where they previously held sway. As a result, the farmers and elephants come into contact, and conflict results: farmers’ fields make wonderful feeding stations for elephants, and elephants make fearsome and sometimes lethal nocturnal crop raiders. The research looked at how elephants and farmers could share the same landscape. The answer turned out to be to control where elephants could go. Over the years, a diverse tool kit of technologies and strategies was developed to keep elephants off farmers’ fields, including using watchtowers, powerful rechargeable torches, loud firecracker sticks, and fences smeared with pungent chili grease and festooned with beehives (both of which elephants hate). Wildlife managers have also built hundreds of kilometers of electric fencing powered by solar panels, with long outriggers to give pushy tuskers a benign shock on their chests. They have employed people to identify individual elephants (for example, from tears in their ears) and to work out which ones have learned to break the fence. They have tranquilized fence breakers and fitted them with radio collars that send their location to the mobile phone network in real time, both to understand how elephants think about risks in the landscape and to offer some kind of early warning. They have tried sawing off the tips of the tusks of fence-breaking elephants (because they use their tusks to short out the wires of fences), and they have tried moving elephants hundreds of kilometers away to areas where it is hoped they will cause less trouble.31 Elephants epitomize the wildness and independence of nature and (with the rest of the “Big 5” wildlife species) carry a whole tourist industry on their backs on the strength of their apparent freedom. Yet, in these settings, they are among the most tightly monitored and controlled animals on earth. Elephant conservationists also make use of multiple other technologies, such as drones, infrared cameras, microphones, helicopters, and satellites, as well as “big data” analysis using artificial intelligence and machine learning to monitor human movement in anti-poaching campaigns. Away from such fraught settings, computers and smartphones are also starting to transform conservation in the fields of public engagement, citizen science, data analysis, and decision support.32

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The Problem of Nature

Similar kinds of innovative and often artificial methods are also being applied in many other dimensions of conservation. One example is the attempt to control wildlife disease. The world’s rarest wild canid, the Ethiopian wolf, is confined to the Bale Mountains of southeastern Ethiopia. It is classified as “endangered” on the IUCN Red List of Threatened Species, with a known (and decreasing) wild population of less than two hundred adults. The disease of rabies is a key threat to the wolf. Rabies also infects local domestic dogs, and conservation efforts have involved both trying to eradicate rabies among dogs (to reduce the disease reservoir) and feeding a vaccine directly to wolves using bait.33 A similar approach to rare-species conservation is being used by the US Fish and Wildlife Service in Montana, which is feeding prairie dogs peanut butter laced with vaccine against sylvatic plague, hoping to boost their population and increase the amount of food available for endangered black-­ footed ferrets, which depend on them; there are even experiments with drones to distribute it.34 The “command and control” approach, mentioned earlier, has long been seen as the key to success in ecological restoration, which has become an increasingly important conservation strategy as the human transformations of the Anthropocene deepen and extend. As an early advocate of restoration put it, the restoration ecologist needs “the ability to restore quickly but to restore at will, controlling, decelerating change as well as accelerating it, reversing it, altering its course, steering it, even preventing it entirely.”35 The scale of plans to restore lost ecologies globally is astonishing. In 2011, the German government and the International Union for Conservation of Nature (IUCN) launched the Bonn Challenge, proposing to restore 3.5 million square kilometers of forest by 2030. Forty-three tropical and subtropical countries (including China, India, and Brazil), have committed nearly 3 million square kilometers of land to this purpose.36 The origins of ecological restoration lie in the reclamation of postindustrial sites and recently transformed ecosystems (such as the lost grasslands and savannah woodlands of the American prairies). Restoration fits on a continuum of ecological management, from the reduction of human



The Problem of Nature 35

impacts, through remediation (of polluted and contaminated sites) and rehabilitation (of areas used for human economic activity or settlement), and ending with ecological restoration.37 The Society for Ecological Restoration (SER) published a handbook in 2004 that defined ecological restoration as “the process of assisting the recovery of an ecosystem that has been degraded, damaged or destroyed.” Classical ecological restoration was focused on the idea of recovery using historical references as goals for restoration projects and emphasized historical fidelity in ecological structure and species composition.38 Ideas have become a little more flexible. In 2016, the United Nations Convention on Biological Diversity added a purpose for restoration, to sustain resilience and conserve biodiversity. The SER now describes the aim of ecological restoration as “to move a degraded ecosystem to a trajectory of recovery that allows adaptation to local and global changes, as well as persistence and evolution of its component species.”39 Conservation scientists recommend that for conservation programs to be successful, specific targets must be set for interventions and projects, and progress must be judged against them. However, as Dutch philosopher Josef Keulartz observes, restoration of past conditions is “a Sisyphean task, akin to paddling upstream into a strong current of global change.” Over longer time periods, it becomes both less easy and less useful to know what state of nature to use as a baseline against which to measure success.40 The ecological past is one of constant change. Over decades, centuries, and millennia, ecosystems have evolved continuously in response to climate, as different sets of species have self-assembled to form ecosystems. Furthermore, it is not easy to put a date on the onset of human impacts. Key moments of intense human action may be obvious in some places. In North or South America or Australia, the arrival of Europeans created a marked historical discontinuity, a step up in the level of destruction of nature that accompanied the colonial annexation of territory. But the first people in these lands had already transformed these landscapes, variously clearing, burning, hunting, grazing domestic animals, and farming. Elsewhere, the history of humanity’s ecological modifications can be hard to disentangle. In Ireland, for example, a combination of deforestation, ag-

36

The Problem of Nature

riculture, and animal husbandry in the Bronze Age (4,500–2,500 years ago) overlapped with (and were perhaps linked to) changes in climate. The history of human transformation of the earth is long and complex: the Anthropocene has deep roots.41 Anthropogenic climate change is a game shifter with respect to human impacts on the earth, and a major driver of ecosystem novelty, as Bill McKibben pointed out very eloquently in his book The End of Nature. He argued that human-caused climate change had, in the twentieth century, brought about the end of nature as something that exists independently of human agency. For practical purposes it is fair enough to say that ecosystems never entered by humans (the bottom of Antarctic sea ice, or scrub on an inaccessible mountaintop) are natural, yet the implication of The End of Nature is that even these places are being changed by anthropogenic climate change.42 Anthropogenic climate change now forms part of the environmental background condition to which all species have to adapt. In any given ecosystem, some species will be lost and others will arrive as climatic conditions change, causing communities to change and reassemble. Some entire ecosystems may change (most obviously on mountains and coasts), while others may undergo subtler shifts as species appear and disappear. Habitat fragmentation will increase the effects, because the ability of species to adapt will be limited by reduced genetic diversity in isolated populations and by the lack of habitat for dispersal or migration. Even in the oceans there are signs of climate-induced ecosystem change. Coral recruitment has decreased in tropical waters, and some reef-building corals are beginning to form “climate refugee” ecosystems in previously coral-­ free areas farther away from the tropics like southern Japan and the Gulf of Mexico. Anthropogenic climate change is creating new ecological trajectories that potentially lie outside historical conditions in all ecosystems on earth.43 Ecological processes are as important in restoration projects as ecosystem structure and composition. As Eric Higgs observes in his book Nature by Design, “No matter how much human agency and intention are applied to the practice of restoration design, natural process kicks in and sometimes takes over completely. What is more, this is typically desirable. Call it wild design.”44



The Problem of Nature 37

Not only does anthropogenic climate change interfere with our idea of naturalness, but so too does the human capacity to create artificial environments. Perhaps the most vividly human-made environments are in cities. And, paradoxically, here many wild species find habitable niches. This is not a new phenomenon: at some stage swallows and swifts moved from rocks and cliffs to house roofs, as some bats did to shingles and barns. Such cohabitations could be seen as a form of unplanned domestication, a one-sided discovery of a usable space where none was intended.45 A whole field of ecology has grown up devoted to the unusual assemblages of species that help form urban ecosystems. In Berlin, for example, ecosystems that had no analogue elsewhere developed on bombed sites after the Second World War. They contained exotic species brought by occupying armies and escaped from gardens, growing on strange substrates. Ecologists in Berlin, themselves isolated due to Cold War politics, developed a wholly new field to analyze these novel, ruderal, and transient ecosystems. Artificial environments and accidentally arriving species make a strange but often valuable pairing in a human-shaped natural world.46 Urban ecosystems emerge under novel and human-created conditions, dominated by human technologies, decisions, and actions. But they also reflect a measure of self-assembly, combining species that have survived human management with those that have hitchhiked on human activities. These emerging ecosystems, lacking an analogue or exact equivalent elsewhere, have begun to attract the attention of ecologists as “novel ecosystems.” They arise from human impacts on animal and plant populations (especially extinctions and introductions); the creation of urban, cultivated, or degraded landscapes; or wider ecological changes (for example, pollution or mining).47 More and more species are “out of place” in the global ecosystem—or rather, have found novel places to be. Naturalization hotspots (measured as the percentage of naturalized aliens in the total flora) include the western and eastern coasts of North America, northwestern Europe, South Africa, southeastern Australia, New Zealand, and India. The New World has almost ten thousand species of naturalized alien plants, and the Old World almost eight thousand.48 Many introduced species become invasive in their new home. In Port-

38

The Problem of Nature

land, Maine, Kent’s partner, Pamela, with Kent and a group of neighbors, have been attempting to revive a five-acre city park. When this effort started, part of Harbor View Park was covered in Japanese knotweed, originally introduced to North America as an ornamental garden plant. It is now on IUCN’s list of the world’s most invasive species, and it had taken over. After they spent two hot, sweaty summers pulling knotweed, it was effectively eliminated. The reward for this labor was the arrival of an even more invasive species, black swallow-wort, a plant with the scary colloquial name of “black dog-strangling vine.” Black swallow-wort had also been introduced to the United States as an ornamental plant (from southwestern Europe) and had escaped. It is a vigorous grower and forms dense patches that suppress all other plants as well as invertebrates. It is harder to get rid of than knotweed, and it is winning the competition to take over the park. Without laborious management, invasive species will drive the struggling native plants into extinction. Harbor View Park would be just mown grass and black swallow-wort, with an occasional crab apple tree poking its head above a mat of vegetation so thick that one might even imagine it strangling the occasional black dog.49 Many alien species become invasive in this way, their aggressive proclivities often revealing themselves long after first arrival. However, others live peaceably in their new homes, staying within gardens or parks or providing ecosystem functions and services, for example, restoring polluted environments. Some may even provide habitat for otherwise threatened species. One example is the Rodrigues fody, a small bird endemic to the smallest of the Mascarene Islands in the Pacific. In the 1960s, its population crashed when forests were cleared for intensive agriculture. It survived not because native woodland was restored, but because it found a new home in plantations of fast-growing non-native trees.50 Nonhuman nature never sits still in the face of the opportunities and threats offered by human activities. Even conservation action is part of this shaping of nature—as the geographer Michael Carolan observes, “conservation policy, once enacted, becomes part of the ecology that it was designed to protect.” Nonhuman life is an actor in its own drama, and natural systems at all scales respond and adapt, with ecosystems taking new forms as they are constituted by new assemblages of species.51 Dramatic examples of nature responding to opportunity are the “ex-



The Problem of Nature 39

clusion zones” created after the Chernobyl nuclear accident in 1986 (in what is now Ukraine and Belarus), and after the Fukushima Daiichi disaster in Japan in 2011. In both cases, a self-assembled ecosystem has developed on abandoned land, a natural “rewilding” experiment. Abandoned by all but a transient human occupation of scientists and, bizarrely, in the case of Chernobyl, postapocalypse tourists, these zones have become de facto nature reserves, colonized by numerous species including a range of large mammals, including wolves and feral dogs at Chernobyl, raccoon dogs at both sites, and Japanese macaques at Fukushima.52 The challenge for conservation is that in many ways these new systems are natural. Their starting point is the result of human action, but the way in which species arrive by themselves is not the result of human choice or the application of human technologies. Unlike the “natural” ecosystems burned, cut, or trapped to maintain nature in the best state for some desired species, these ecosystems are self-assembled or, in the language often used in the United Kingdom, “self-willed.” This curious phrase reflects the way that they have formed through natural responses to ecological opportunity, even if the conditions starting their development are human-made. For conservation, the loss of the biodiversity that evolved in the pre­ human world is a deep-seated and fundamental concern. Such loss has cultural significance in the same way that the loss of ancient works of art or significant historical buildings is important (Notre Dame in Paris, for example). It is the apparent lack of human influences in the famous wild spaces of the earth (the Congo, the Pantanal, Okavango Delta, the high Himalaya, or the Arctic) that makes them so wonderful to the nature lover or conservationist. The thing that gives cultural value to nature is its seeming freedom from human artifice. Yet even human-influenced nature can have an essential naturalness. The British ecologist George Peterken usefully distinguishes between different categories of naturalness. He contrasted “past naturalness” (the state of nature before humans arrived), with “future naturalness” (the state of nature in the future if all deliberate human management is stopped). Even a much-transformed human system can be “natural” in this “future-natural” sense, even if it has no analogue in the past.53

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The Problem of Nature

Ecosystems are not stable and unchanging, but constantly adjusting to natural processes operating over different scales in space and time. In the Anthropocene, human actions (including conservation’s intensive management of ecosystems and populations) weave themselves into the ecology of places. What humans call nature is therefore a hybrid of biological and cultural influences, an amalgam of human and nonhuman actions— what Emma Marris called “a rambunctious garden.” This idea is challenging, because it suggests that the things conservationists seek to protect are partly (and increasingly) “artificial,” at least in part. Accepting this opens up questions about the value of conserving ecosystems that have been strongly shaped by human action or have self-assembled under human-­ dominated conditions.54 The Society for Ecological Restoration notes in its Primer that restoration can lead to unexpected results. One example is when actions designed to help the recovery of a native species also stimulate other changes, for example, allowing weedy plants in soil seed banks to flourish. The resulting ecosystems, self-organizing and adapting to circumstances, may be different from those once on the site, but they can be valuable and self-sustaining. A measure of openness and flexibility may be required in the face of natural ecosystem dynamics.55 The best examples of attempts to create a “future-natural” state come from the rewilding movement, which dates back to the 1990s. In 1998, Michael Soulé and Reed Noss published a paper arguing that without top carnivores, such as cougars, jaguars, wolves, wolverines, grizzlies, or black bears, North America’s native nature seemed “somehow incomplete, truncated, overly tame.” They called for the creation of large areas of “wilderness” in North America, within which large predators could re­ establish former ecological processes.56 In 2005, a paper in the journal Nature called for “Pleistocene re-wilding,” making the case that the ecology of North America and Eurasia had been shaped by large mammals that had become extinct at the end of the Ice Age, and that these species (or modern-day substitutes) should be reintroduced to restart dynamic ecological processes. Segey Zimov, architect of a Pleistocene “rewilding” project in Siberia, argues that the intro-



The Problem of Nature 41

duction of modern large herbivores could restore the complex ecology of former Arctic grasslands as well as increase carbon sequestration.57 In a number of countries, conservationists have introduced predators (for example, wolves in Yellowstone National Park, or lynx in Europe), or “keystone” herbivores, such as beavers, in the hope of restarting “tropic cascades” of predators and prey. In Yellowstone, the interaction of elk and introduced beavers and wolves transformed the recruitment of quaking aspen, transforming riparian ecosystems. In Europe, there has also been widespread experimentation with traditional breeds of domesticated herbivores (for example, Konik ponies from Poland, or hairy Highland cattle from Scotland) to simulate the grazing and landscape disturbance that once shaped ecologies.58 As rewilding practices mature, a theoretical framework is emerging, based on the maintenance of complexity and resilience through interactions among trophic (predator-prey) relationships, natural disturbances, and dispersal. While rewilding project managers may set specific targets (for example, establishment of a self-sustaining population of a locally extinct predator) and be able to state the desired ecological effects of such an introduction (for example, the generation of a trophic cascade or a new pattern of ecosystems), the details of specific ecological outcomes are hard to predict accurately. One review describes the effects of rewilding as likely to be “indirect and unexpected.” Engagement with rewilding reflects an acceptance of open-endedness.59 Some ecologists criticize the term “rewilding” as merely a buzzword and argue that it is indistinguishable from existing terms such as “restoration” or “translocation.” Such critics are correct that the word is used to attract the attention of policy makers or to attract funds. Yet the concept is useful, as a subfield of restoration that promotes the restoration of selfsustaining and complex ecosystems through the natural ecological processes involved in population, community, and ecosystem dynamics.60 One trial of this idea is at the Wicken Fen Vision, a British nature reserve in eastern England, where Bill’s partner, Francine, has done research as an ecologist for many years. Here the National Trust (a national UK conservation charity) owns a small fragment of the old fenland, a wetland that once stretched from the city of Cambridge to the sea. The old Wicken

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The Problem of Nature

Fen National Nature Reserve covers an area of just 170 hectares but has extraordinary biodiversity (nine thousand recorded species). It is intensively managed to maintain existing biodiversity using rotational cutting and vegetation removal and the maintenance of water tables above the ground level of the surrounding reclaimed and drained farmland.61 In 1999 the National Trust announced a “100 year vision” to create over three thousand hectares of wetland out of drained and intensively farmed arable land around the old reserve. It hoped that this new area would contribute to the survival of some of the rare plant and invertebrate species on the old fen. However, the trust recognized that it would not be technically or economically feasible to replicate historical habitats, because drainage and intensive arable farming had created a novel starting point for ecosystem restoration. There were no analogues to inform expectations of ecological outcomes at the site. The project therefore ­adopted an open-ended approach and set a goal of allowing or establishing ecological processes (natural regeneration, naturalistic grazing, and fluctuating shallow water tables) on a large scale. Secondary goals were to create a dynamic wetland landscape characterized by mobile vegetation mosaics and to increase ecosystem service provision. No specific species or habitat targets were set: it was accepted that species assemblages would be novel, containing mixtures of native fen-specific species, native generalists (both wetland and terrestrial species), arable-specific species, and non-­ native species.62 To date, the project covers about six hundred hectares. Drainage ditches have been blocked to enable water tables to rise and fluctuate, and semiferal grazing animals (Konik ponies and Highland cattle) serve as agents of ecosystem disturbance. A habitat mosaic of shallow water bodies, reedbeds, wet grassland, dry grassland, and scrub has developed. The reserve has large numbers of wintering wildfowl and supports rare breeding marsh harriers, bitterns, and cranes. In whatever way it is idealized, nature does not sit still in the face of ever-increasing human pressure and influence. It interacts with and responds to human and nonhuman influences. It has done so in the past, does so in the present, and will do so in the future. A metric of “natu­ ralness” is the capacity of nonhuman lives and systems to live and act or



The Problem of Nature 43

“perform” according to their own dynamic ways that are not dependent on human wishes and control. Conservation faces a tricky paradox. To protect existing ecosystem diversity and species, it is sometimes necessary to use highly artificial methods. Preservation of the natural increasingly depends on markedly unnatural interventions. Yet organisms spontaneously respond and adapt to artificial starting points, resulting in the emergence of novel combinations of species that may have no analogue elsewhere. Such responses are “natural.” The ecosystems that arise are natural, but also novel. It is a serious thing to abandon the simplistic category of “natural” as a basis for conservation thinking, to address the dilemma that the preservation of naturalness often demands extreme artificial methods, while new ecosystems that are emerging do not resemble those of any known past. As the geographer Jamie Lorimer observes, “When one can no longer make recourse to Nature, what forms and trajectories of difference matter? Who decides? On what grounds? And through what processes?” There will be difficult decisions and difficult arguments about what forms of social nature are permitted.63 Uncertainty about how to think about novel and hybrid ecosystems makes it hard for conservation to come to terms with functioning ecosystems with novel elements and starting points, and with ecosystems that function but lack the full complement of the species thought once to exist there. But, as Rachel Carson pointed out, “Like the resource it seeks to protect, wildlife conservation must be dynamic, changing as conditions change, seeking always to become more effective.”64 Ecological novelty is a feature of ecosystems and of the species within them. Such novelty may be the result of human action but may also result from ecological interactions or changing environmental conditions. Such circumstances are novel if they lie outside the range of conditions experienced by a species during its evolutionary history (biotic factors, past interactions with competitors, predators, prey, or parasites). Selection pressures associated with novel conditions shape the morphological traits, behavior, and fitness of all organisms.65 So far, most debates about naturalness and conservation have taken place at the level of the species and ecosystem. These are complex enough, but debate has at least started. But the question of naturalness at the ge-

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The Problem of Nature

nomic level is also important. This is still terra incognita for conservation. What might it mean to talk about the naturalness of the genetic makeup of an organism? In chapter 3, we start to explore these questions by thinking through biological diversity at the level of the gene.

3 Nature’s Diversity

When Kent went looking for giant anteaters in the wild in Brazil, he was told to look for “an animal that was too long.” Sure enough, the animal itself, a delightfully odd-looking beast the size of a German shepherd dog, with a very long nose and an even longer, and very bushy, tail, showed how accurate that description was. In Portuguese the giant anteater is called tamanduá bandeira—“tamanduá” from the Indigenous Tupi language meaning “ant hunter,” and “bandeira” from the Portuguese for “flag.” They don’t see very well, but their noses give them an unerring ability to locate termites in their mounds, which rise like hard mud sandcastles from the surrounding grasslands. The giant anteater’s diet of termites and ants is incredibly specialized. Not only are its body and even skeleton adapted to this particular diet, but so too is everything else about it. Its front feet are shod with long claws that serve as powerful tools to break through the hardened mud walls or termitaria. Inside the anteater’s long snout, bereft of teeth, is an even longer tongue covered in sticky saliva, to pull termites from deep within their tunnels. It turns out that the giant anteater is a remarkably fast eater, swallowing termites and squashing them in its mouth but also in a specially toughened stomach. The giant anteater was described by the great Swedish taxonomist Carl

Giant anteater 45

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von Linné (Carl Linnaeus) in 1758. Named as Myrmecophaga tridactyla (literally, “three toed ant-eater”), it is placed with the other South American anteaters in the suborder Vermilingua (literally, with apologies to The Lord of the Rings fans, “wormtongues”). Anteaters are related to sloths, and with them to armadillos. Kent’s study animals were living their anteater lives among termite mounds in Emas National Park in central-western Brazil. This wonderful and still-little-known park protects some of the best examples of the cerrado biome. Cerrado occurs to the south of the rain forests of the Amazon and covers a huge area (a fifth of Brazil). It is dry, with thirteen hundred millimeters of rainfall a year, falling between September and March, supporting a landscape of forest and open grassland savanna. The Parque Nacional das Emas is both beautiful and diverse. Habitats range from open grassland dotted with termite mounds, through dry tropical forest, to flooded grasslands with their hidden frogs waiting until the rain. Along the rivers are strips of cool forest, where the sweat and dust of a day in the sun can be washed off in the fast-flowing clear water. Giant anteaters are everywhere in the park, sleeping on low termite mounds, carrying their babies on their backs, and incessantly feeding on ants and termites. Brazil has a lot of termites: about 150 species, half of them found in the cerrado. There are big ones with strong pincers, small speedy ones that spray chemicals from their nozzle-shaped heads, and slow gray ones marbled with the soil they consume. Some species build mounds up to two meters tall, and these serve as home to their builders as well as over 20 other species of termites, over 20 species of ants, and countless cockroaches, beetles, spiders, millipedes, and centipedes. There is even one beetle, only described in the last few decades, whose larvae bioluminesce at night, turning the many mounds where they are found into fairy castles of lights.1 The cerrado is a natural wonder, but it is shrinking. In 1960, the Brazilian government chose to locate the remarkable modernist architecture of their new federal capital, Brasília, in the heart of the cerrado, and the area began to be heavily promoted for agricultural development. Rapid growth in the Brazilian economy in the 1960s and 1970s was accompanied by large-scale investment in the rural sector and rapid land clearance, both in Amazonia and in the savanna of the cerrado farther south. Initially land



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was cleared for cattle ranching, but from the 1990s, a boom in the mechanized production of soy (soya bean) became the major driver of land-cover change. Transnational agribusiness companies began to invest in large-­ scale soy production for export, driven by the ever-increasing market in China for animal feed. Genetically modified seed resistant to glyphosate-­ based herbicides (such as Monsanto’s Roundup) was approved in Argentina in 1995 and smuggled into Brazil before being approved in 2003. By 2015, Brazil was the world’s second largest producer of soy, with a third of global production.2 As soy has expanded, the cerrado and its biodiversity has retreated, leaving remaining patches of unploughed savanna and their natural riches as tattered patches in a backcloth of modern agriculture. The cerrado is in serious trouble, with deforestation rates higher than the Amazon rain forest. It fits the familiar conservation story line of wild nature shrinking in the face of human demand for land. Emas National Park is an island in this landscape, though admittedly a big one. The species diversity of the cerrado is remarkable: it is the largest, oldest, and biologically richest savanna ecosystem in the world. It is one of the most diverse of all tropical habitats, with over 1,200 terrestrial vertebrates and over 11,000 species of plants. It sustains between 350 and 400 species of vascular plants per hectare and populations of giant anteater, giant armadillo, maned wolf, jaguar, rhea, the blue-and-yellow macaw, and the buriti palm. Many species are endemic, meaning that they do not occur anywhere else on earth, only in the cerrado. There is also an incredible diversity of microorganisms, bacteria, fungi, and viruses, forming their own mini-ecosystems (microbiomes) in the guts of termites, the soil of the grasslands, the skin of a streamer-tailed tyrant—everywhere you look.3 The remarkable wildlife and ecosystems of the cerrado are in one sense unique, in that no other region of the world (even those with a similar dry seasonal climate and savanna vegetation) has the same mix of species, many of which are unique to these South American landscapes. But the cerrado ecosystem shares one characteristic with other ecosystems across the world that have been little changed by human activity—its diversity. It holds an astonishing number of different life-forms across all taxa, from mammals to insects to plants to bacteria and fungi—and, of course, to termites.

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It was the conservation significance of this ecosystem—a mix of the overall number of species and the number that were endemic, found nowhere else—that led the US-based organization Conservation International to identify the cerrado as one of the world’s “biodiversity hotspots,” one of thirty-six regions across the world that together represent 2.4 percent of the earth’s land surface but support more than half of the world’s endemic plant species and nearly 43 percent of bird, mammal, reptile, and amphibian species. The cerrado’s conservation value was also the reason why the Brazilian government created the Emas National Park in 1961, and it became a UNESCO World Heritage site in 2001.4 Why is it such an important place for the global conservation community as well as the citizens of Brazil? Emas, like many other protected areas around the world, managed by governments, private citizens, businesses, churches, and Indigenous peoples, is an essential component in the global effort to conserve biodiversity, in all its richness, across all three of its manifestations—genes, species, and ecosystems. Emas contains a globally significant portfolio of all three: a diversity of ecosystems, from open grassland to riverine forest; a diversity of species—just think of the termites; and a diversity of genes, a veritable landscape of genes in the plants, animals, and microbes. Biological diversity is essential to proper functioning of ecosystems. As we have seen, the diversity of species (and in particular the existence of endemic and rare species bigger than a warbler or tarantula) is of huge importance for conservationists. The existence and variety of life-forms evoke human wonder, as the popularity of successive blockbuster television documentary series attests. The diversity of species includes many that have proved essential to human survival and economy on earth (from food crops and animals or the fish in the sea to penicillin or the chef’s spice rack). As far as astronomers know, earth’s life and the diversity of its life make it unique in the universe. Preventing extinction and ensuring the continuity of diversity in all its forms are purposes of much of conservation’s efforts and the reasons for its popular support. Where did the variety of life come from? The answer, as every biology student is taught in school, is evolution. But how did diversity evolve? The simple and traditional answer is slowly—mostly very slowly. Some-



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time, about four billion years ago, life emerged on earth, preserved in ancient rocks in Greenland and Western Australia. The putative first organism has been given the unsexy name of LUCA—standing for Last Universal Common Ancestor. It is from LUCA that all of the incredible diversity of life on earth is thought to have sprung. The first organisms were not very exciting—single-celled archaea and bacteria forming microbial mats in the oceans that dominated the earth under an atmosphere hostile to almost all modern forms of life. About 3.5 million years ago, living organisms began to capture and use the sun’s energy for the first time. In turn, they began to churn out oxygen and transform the earth’s atmosphere. Just under 2 million years ago, more complex single-celled organisms began to emerge, which had structures within their cells. Just over 1.5 billion years ago, multicelled organisms, in which different cells performed specialized functions, appeared in the fossil record. Complex plants had emerged by 850 million years ago, and about half a million years ago what geologists call the Phanerozoic era had begun. It saw the successive emergence of all the forms of complex life we know today: fish, vascular plants, amphibians, reptiles, and trees.5 The first mammals and the first dinosaurs appeared a quarter of a million years ago, followed by birds and flowering plants. The earth had become the world that’s now familiar from the film Jurassic Park or the TV show Walking with Dinosaurs. The end of the Cretaceous period (65 million years ago) saw the sudden extinction of the dinosaurs. Mammalian diversity exploded once the dinosaurs were gone. Very recently, about 2 million years ago, the species we have come to call “human” began to emerge.6 Diversity can be identified in the strangest forms: tube worms living near thermal vents among the benthos in the pitch dark and high pressure of the ocean’s seafloor; bacteria living under a kilometer of ice in an Antarctic lake; marine worms with no mouth and no gut that eat whale carcasses; English oak trees living up to a thousand years; the ten-centimeters-long pink fairy armadillo that lives underground in the dry lands of Argentina; and the bowerbirds of New Guinean forests that build and decorate elaborate structures to court choosy females. There are estimated to be between 2.2 million and 3.8 million species of fungus across the world, and the atmosphere itself is alive with microbial diversity that rivals that of any

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other ecosystem. In one study, hundreds of millions of viruses were found in one square meter of ground. Diversity seems to abound at whatever scale you look, but particularly at the microscopic scale.7 For every species that currently exists, others have gone extinct at some point in geological time. So in South America, cerrado vegetation has existed over tens of thousands of years on the central Brazilian plateau, on ancient soils from which nutrients have long been leached away. On it once roamed a bestiary of animals, some from the same taxonomic lineages as today’s inhabitant. Flightless birds (dubbed “giant terror birds”) up to three meters tall were the top predators in South America in the Cenozoic era (62 million to 1.8 million years ago), while glyptodonts, armadillos the size of a small car, roamed the plateau, and there were ground sloths the size of bears. All of these species are long gone, extinct like so many others. Despite the many extinctions that have taken place, the earth is a species-rich planet. No one knows how many species there are on earth, though a 2019 UN report stated that the consensus number is 8 million (including 5.5 million insects), but not including microbes. In 2018, at least 229 new species were described, including 120 wasps, 34 sea slugs, 28 ants, 19 fish, 7 flowering plants, 7 spiders, 4 eels, 3 sharks, 2 water bears, 1 frog, 1 snake, 1 seahorse, 1 moss, and 1 liverwort.8 This variety of life has long fascinated naturalists. It awoke in humans an insatiable desire to name and classify based on a wide variety of characteristics of organisms, from smell to color to spiritual qualities. In the Western world this led to systems of organization and classification to catalog life’s variety in museums and herbariums full of preserved specimens, and in time to zoos and botanic gardens, to trophy collecting by hunters, and eventually to the checklists of obsessive birders, to iNaturalist.org, and today’s photographic websites where every new sighting is recorded. The early collections revealed the diversity of tropical regions to European philosophers and naturalists and further fed this hunger. The specimens and the stories their collectors brought back from tropical regions not only stimulated the centuries-long land grab of imperialism, but also had a huge impact on ideas about the earth and the diversity of life upon it. In the eighteenth and nineteenth centuries, gentlemen built cabinets of curiosities, collections of natural history, rooms stuffed with the obscure, the arcane, and the exotic—one of each type. The glass case full of



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stuffed hummingbirds became a commonplace of the wealthy Victorian drawing room. Museums and herbaria documented the many, now-named, types of animals and plants, and zoological and botanical gardens made them available for public wonder. At the heart of this was the urge to describe and document the myriad kinds of life. Working in the Swedish university city of Uppsala in the eighteenth century, Carl von Linné, the father of modern scientific taxonomy, published Systema Naturae in 1735, marking the beginning of biology’s fascination with species. The system of scientific classification based on species became the currency for modern taxonomic science and the foundation of much of conservation’s efforts. Characterizing the different kinds of organisms became an obsession in Europe and America with the work of naturalists formalized into the science of taxonomy. In museum cabinets in Paris, London, Berlin, and Washington, type specimens were laid out, each tagged to the place where they were shot or snipped from the branch, and the name of the person who first described them. Each species was separated by a distinguishing physical feature, sometimes obvious (like plumage on a bird), sometimes hidden (the bones in a sea lion’s ear). Affinities between species and among sets of species were explored, and a hierarchical system of classification was teased out on the basis of physical similarity: species, genus, family, order, class, phylum, and kingdom. Scientists needed a way to standardize their classifications. A binomial system of scientific nomenclature was adopted, using that lingua franca of the educated European class, Latin and Greek: a second name that referred to the unique species, and a first name that referred to the genus (the set of similar species) within a given family of organisms. Thus, the maned wolf of the cerrado, is given the name Chrysocyon brachyurus. It is called a wolf and looks like a very long-legged golden dog, but it is neither. Taxonomists place it in a genus all of its own within the wider dog family (Canidae). The names of different natural types became a problem. Many were well known by colloquial names to people who fished, hunted, or collected medicinal herbs. The United States, for example, boasts the world’s highest diversity of freshwater mussels, and they acquired wonderful names from European settlers, like orangefoot pimpleback, sad elliptio, and rough

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fatmucket. But such names did not mean the same things in all places, and the habit of European settlers of assigning to new animals or plants the names of those from the mother country created extra confusion. The American robin, for example, is a very different bird from the European robin, after which it was nostalgically named. The traditional basis of taxonomy is physical description. Specimens are still killed and returned to herbaria and museums, to be pored over by experts and named, their lineages and affiliations teased out and published. Naming has always been quirky. Many species were named after the person who described them, or after a patron. More recently, an attempt has been made to make names descriptive, although naming can also be somewhat eccentric: in 2019 an Australian jumping spider was called Jotus karllagerfeldi, because its dark eyes and black-and-white-striped legs reminded the taxonomists of the couturier Karl Lagerfeld.9 However, the advent of faster and better technologies for reading genes and genomes is opening a new chapter in taxonomy, separating species that cannot necessarily be differentiated physically, and certainly not visually. Such reworking of well-established taxonomies using genetic differences is becoming more common—for example, mouse lemurs were described as one species in 1931, while genetic analysis has led to a current listing of eighteen species, or twenty-one: different kinds of genetic tests yield conflicting results.10 Two nineteenth-century Englishmen, Charles Darwin and Alfred Wallace shaped the way we think about natural diversity today. These two had the greatest influence on the way natural diversity is understood today. Although they worked in very different regions, they both asked the same question: How is it possible to explain the variety of life on earth? People have long known that all life is not the same. Of course, mammoths are different from proboscis monkeys or yew trees from eelgrass, but not even all sheep are the same. There are differences of shape, size, speed of growth, length of wool, and ability to withstand drought. The same goes for humans: think about skin color, or hair, that fickle characteristic of perceived beauty; of course, we do not all look the same. The difference between individuals, the variety of life at the finest scale, has been part and parcel of every human observer’s experience. The variety within a single nonhuman species was already widely recog-



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nized by the middle of the nineteenth century, not least by one of Darwin’s mentors at Cambridge, John Henslow, founder of the university’s botanical garden. He assembled an herbarium of the British flora to show variation within species and to determine the limits between species. He was a religious man who believed the orthodoxy of the day, that species were stable and did not evolve. Darwin, whom Henslow recommended to the captain of HMS Beagle, was to show something different.11 People understood that offspring tended to look like their parents and that there was some factor that caused this to happen. This factor took many forms in different cultures. In the first century BC, Pythagoras, a Greek scholar, advanced the notion that hereditary information was carried in male semen, which collected instructions to transmit to offspring by absorbing vapors from each of the individual parts, and then, when transferred to the mother’s womb, matured into a fetus. And Darwin himself posited the existence of particles he called “gemmules,” which somehow passed information to sperm and egg and then to offspring.12 In whatever way the mechanism was explained, people understood and worked with variation in domesticating wild species. Over generations and generations, our ancestors turned species of wild grass into wheat, corn, rice, sorghum, millet, or the Ethiopian cereal teff. They tamed wild aurochs and turned them into a whole bestiary of domestic cattle breeds, and they learned to take wild yeast and turn it into something that could be relied on to make bread rise and to ferment beer to celebrate the solstice. But there was little understanding of what caused the variety between individuals and how the distinctive characteristics of animals or plants were passed on through generations. Both Wallace and Darwin figured out the answer to the question of how variation was generated. They worked independently, Wallace in the Malay Archipelago, and Darwin in South America and the Galápagos Islands, on the famous voyage of HMS Beagle between 1831 and 1836, and later back in the United Kingdom. By 1858, Wallace had worked out that the fittest individuals survived and reproduced, passing their advantageous characteristics on to their offspring. In that year, he wrote to Darwin enclosing a paper (“On the Tendency of Varieties to Depart Indefinitely from the Original Type”) setting out his ideas. At the time, Darwin, was himself laboring over his own paper, drawing together his reflections

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on natural varieties and selection, stemming from observations made on the Beagle expedition two decades before. In the end, they jointly published a paper entitled “On the Tendency of Species to Form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection.” It was read to the Linnean Society on July 1, 1858, introduced by no lesser luminaries than Charles Lyell and Joseph Hooker, before being published a few months later in the Journal of the Proceedings of the Linnean Society: Zoology. What they wrote provided the framework for evolution as understood today, and eventually for the science of genetics. Their argument is best known now through Darwin’s book On the Origin of Species (1859) in which he set out the famous thesis about how the power of natural selection, working on the innate variety between individuals, could drive the evolution of traits. The full title does a good job of explaining its purpose and argument: On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life.13 Darwin thought of natural selection in terms of the selection of traits that are advantageous, in just the same way as a farmer would select the best characteristics of a water buffalo or a variety of potato. He wrote, “If variations useful to any organic being do occur, assuredly individuals thus characterized will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterized. This principle of preservation, I have called, for the sake of brevity, Natural Selection.” Darwin took the term “selection” from the selection practiced by animal and plant breeders.14 Though Darwin’s name is always attached to the concept of evolution, he did not use that term in his book, referring instead to “transmutation.” The word “evolution” appears to have entered English from the Greek via Latin and was used to refer to a military maneuver. In classical Latin, however, the word denoted the unrolling of a scroll, and it was so used in English in the seventeenth century. The term was used in the 1830s by the famous geologist Charles Lyell to refer to the process of development— the meaning we now recognize in evolution.15 Darwin and the many biologists who followed him saw that there was something that both gave organisms their individual characteristics (explaining why two species were so different and no two individuals were



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alike) and that could be passed from one generation to the next (because the offspring of giant anteaters are still giant anteaters, not jaguars or armadillos). But it took a long time to work out how this worked. Only at the end of the nineteenth century did a concept emerge to describe that basic unit of natural variety and inheritance that was responsible for this process. That concept was the gene—and it would come to be understood as the unit of heredity through which evolution could take place. Evolution is responsible for the incredible diversity of life on earth. It is a process whose results, the naturally evolved diversity of life, are the conservationist’s passion. Darwin explained the power of natural selection and in turn laid the groundwork for our understanding of evolution. But he did not know what caused the variety between individuals. Indeed, the extraordinary thing about Darwin and Wallace and their novel idea of evolution was that they had no notion of any mechanism by which what they proposed would work. Evidence for this mechanism did exist, generated in a highly unlikely place—a monastery in what is now the Czech Republic. Gregor Mendel, born in 1822, was a failed teacher, a depressive, and an Augustinian friar with a passion for botany. He set out to understand the ways in which variation in plants was passed through generations. Mendel chose the edible pea as his experimental plant because its different varieties were well known, the plants were easy to grow, and their pollination could be well controlled. He chose seven “traits,” or characteristics, of pea plants and seeds, including seed color, seed form, and plant height and bred strains whose offspring were the same for the key trait. Mendel then crossed varieties (for example, round versus wrinkled peas, short versus tall plants). By repeatedly crossing different varieties over generations and years and keeping painstaking records of the results of all these crosses, he was able to establish the basic rules of inheritance. He showed that the traits he was using did not “blend,” but, instead, offspring had parental traits in entirely predictable proportions. He hypothesized that there were two units of heredity relating to each trait that were inherited independently from each other, one from each parent. He showed that a set of laws governed the ways this inheritance took place. He had effectively deduced the existence of the “gene,” but, of course, he didn’t know the word, instead calling these units “atoms of inheritance.” Unfortunately for Mendel he published the results of his painstaking

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work in the proceedings of the local society of naturalists in Brno (Brünn), now in the Czech Republic, where it went more or less unnoticed. It was not until 1900, sixteen years after Mendel’s death, that his work was rediscovered by Hugo de Vries, Carl Correns, and Erich von Tschermak, three scientists working independently and publishing in the same year. Mendel’s results, which came to be seen as “laws,” found a receptive audience in the scientific community of Europe, the United Kingdom, and the United States and formed a key part of the new field called “genetics.” In 1906 the Englishman William Bateson used the term “genetics” to describe a science “devoted to the elucidation of the phenomena of heredity and variation”: in other words, to the physiology of descent. Decades of work followed, trying to elucidate what genes were, in biochemical terms, and how the machinery behind heredity based upon them actually worked.16 Genes are made up of a complex molecule called deoxyribonucleic acid, or DNA, was first discovered in the late 1860s by Swiss chemist Friedrich Miescher. In 1869, he identified what he called “nuclein” inside the nuclei of human white blood cells, later renaming it “nucleic acid” and then “deoxyribonucleic acid,” or DNA. But the chemical structure of DNA remained a mystery for many long decades. The key (and Nobel Prize– winning) breakthrough was by James Watson and Francis Crick in 1953 (apocryphally in a Cambridge pub), although they drew on the X-ray crystallography of Rosalind Franklin and her supervisor, Maurice Wilkins, to do it. The clever thing that Watson and Crick did was to show that DNA was a two-stranded molecule twisted into the shape of a double helix, a pair of twisted ladders coiled around each other.17 In chemical terms, DNA is made up of building blocks called nucleotides (the “N” in DNA). There are just four of them in DNA: adenine, cytosine, guanine, and thymine. These four nucleotides each contain a sugar (deoxyribose—the “D” in DNA) and a phosphate group (the acid, or the “A”). Two nucleotides form the “rungs” in the winding ladder of the DNA molecule, one reaching out from each side of the ladder, joined in the middle by a hydrogen bond. The sidepieces of the ladder are formed by the sugar and phosphate molecules. A nucleotide always pairs together with a complementary partner on the opposite side, forming



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what is called a “base pair”: adenine always pairs with thymine, and cytosine pairs with guanine. DNA can be thought of as written in an alphabet of four letters, A, C, G, and T, one for each nucleotide. The simplicity of the elements within the DNA molecule is deceptive. Despite the limited number of components (only the four letters), the chemical variety represented in the millions of pairs of nucleotides spiraling up in coils of DNA can represent an almost infinite variety of life. For example, the DNA of domesticated barley contains five billion base pairs.18 DNA exists in long strands that spend most of their time tightly coiled inside the nucleus of cells. The coiling of DNA is an absolute necessity because the length of DNA strands is immense compared to the tiny dimensions of cells. There is no less than one meter of DNA in a single human cell nucleus, coiled into a space just six microns across.19 In eukaryotes—multicelled organisms, including humans, houseplants, fish, and fleas, with cells containing membrane-bound organelles, such as a nucleus—DNA is found not only in the nucleus, but also in multiple organelles such as mitochondria, where energy production occurs. In prokaryotes—single-celled organisms including bacteria and archaea, without a distinct nucleus bound by a membrane—DNA occurs in a single, circular, double-stranded DNA molecule. Eukaryotes have orders of magnitude more DNA than prokaryotes. Viruses, a third category of life, being neither prokaryotes nor eukaryotes, may have some DNA, but not very much. DNA has several vital functions. First and foremost, it stores genetic information. The sequence of nucleotides in a cell’s DNA is distinct between individuals, species, and (in organisms that reproduce sexually) normally unique. DNA copies that information into future generations of cells, to make sure that new cells are the same as the old. And it expresses that information in the form of proteins that go on to build the organism’s structure and enable it to function. DNA can be thought of as the “software” that programs the hardware of the cell and, hence, of the organism. DNA’s ability to replicate itself is a critical feature of life, because as organisms develop or grow, cells have to make new cells. Each new cell needs to be an exact copy of the original cell and to carry an exact copy of the original DNA. The copying process is done by another kind of nucleic acid, ribonucleic acid, or RNA. RNA is a bit like DNA but has a

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simpler structure. Unlike DNA, it is single-stranded, composed of ribose sugar, four nucleotide bases, and a phosphate. The replication of DNA involves unzipping the hydrogen bonds in the DNA strand, and each single “unzipped” chain acts as a template for the assembly of a new doublehelix DNA chain, with one strand from the original and one created anew. This process is astonishingly fast: 33 nucleotides are added per second in eukaryotes, and an astonishing 833 per second in prokaryotes. But even this is slow when a strand of DNA might be 200 million base pairs long, so replication happens simultaneously at multiple points along the length of the DNA strand.20 Other functions are performed by different types of RNA. Most importantly, they carry out the molecular messages from DNA by coding, decoding, regulating, and affecting the expression of genes, but recent work has shown that RNA has a variety of other functions, for example, serving as an enzyme to speed up chemical reactions.21 Geneticists have identified particular lengths of DNA that are responsible for specific effects. These sequences are what we refer to when we speak of “genes.” Genes are not discrete physical units, but simply a name given to lengths of DNA identified as having particular functions. Each gene consists of a defined sequence of bases. Genes can be thought of as a code—like the lines of code that comprise computer software—that directs the biological machinery of the cell, telling it what to do and when to do it. Different genes are devoted to different functions that are vital for the cell, such as making a cell wall, keeping unwanted substances out of the cell, and repairing damage. Geneticists talk about the characteristics or attributes of an organism that are expressed by genes as “traits.” They might include physical properties (for example, the color of a bird’s plumage, color of a human eye, or the shape of a tree’s leaf) and behavioral characteristics (for example, time of bud-burst in spring or a bird’s instinct to migrate south in autumn). The set of genes that is responsible for a particular trait is referred to as the “genotype.” The rise of tools that have allowed humans to peer into and begin to understand DNA has focused attention on genes. This process has sometimes led to the simplistic, and incorrect, assumption that the diversity of life is only shaped by genes. In fact, an organism cannot be reduced to its



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genetic code, but instead is a complicated mix of genetics, development, and ecology. Additionally, traits are often coded by more than one gene, and, equally, single genes may also influence more than one trait. The links between genetic code and the appearance of an organism are multiple, subtle, and complex. The characteristics of an organism are determined not only by the deterministic coding of its genes but also by the influence of environmental conditions. In 1911, a Danish scientist, Wilhelm Johannsen, coined the term “phenotype” for the physical expression, or characteristics, of that trait. The phenotype of an organism is the physical manifestation of genes plus the influences of the environment. For example, all flamingo eggs that hatch will produce flamingos because that’s what flamingo genes are programmed to do—that is their genotype. But not all flamingos are equally pink. Their color comes from pigments in their diet of brine shrimp and blue-green algae. A bright pink flamingo is therefore the result of its genes plus its diet, and it is possible to raise a white flamingo if the birds don’t get these pigments. Biology students are conventionally taught that DNA instructs RNA, and RNA builds proteins using combinations of twenty amino acids in specific sequences. Different genes work to construct different proteins. The order in which amino acids come determines the protein’s structure (for they are folded in unique shapes) and their function in the organism. In eukaryotes, proteins are involved in a huge range of structures and in processes within and between cells, from signaling, transportation, and storage, to the control of other processes (enzymes are proteins, for example) and fighting infection (antibodies in blood are also proteins). In the coils of DNA, protein-constructing genes are interspersed with other sequences of bases (called “regulatory sequences”) that turn these genes on and off. The unique sequence of bases in each gene creates the instructions that dictate what proteins are made and when, and therefore how the cell functions. By the end of the twentieth century, research into how genetic material works in different forms of life had made clear that this standard explanation of what genes are and do was only a first approximation. When botanist Wilhelm Johannsen introduced the term “gene” in 1905, he was attempting to put a name on an unidentified object that determined the

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traits of developing organisms. Yet the gene has remained a somewhat abstract concept, hard to tie to the physical scaffolding of the DNA molecule. In his book The Gene: An Intimate History, Siddhartha Mukherjee observes that the gene was “created to mark a function; it was an abstraction. A gene was defined by what a gene does; it was a carrier of hereditary information.”22 Scientists have struggled with how exactly to define a gene, and Rheinberger and Müller-Wille argue that, despite the widely accepted definition of the gene we have provided above, “a simple and universally accepted definition of the gene never existed.” Nevertheless, the gene is an important concept for explaining how proteins are formed and how organisms function, particularly in the case of microbes; for plants and animals it seems to be much more complicated.23 The ways that a gene is expressed is not straightforward. This means that finding a gene that controls something you care about, say disease resistance or drought tolerance, is not simple. So, to take an example from conservation work, a great deal of effort has gone into the conservation of the California condor. Condors almost became extinct in the twentieth century through loss of habitat and food, shooting, and poisoning from lead shot ingested when feeding on dead animals. In 1967 they were listed as endangered, and in 1982 the US Fish and Wildlife Service began a captive breeding program. By 1984 there were only 14 individuals in the world, and all were taken into captivity. The program has had some success, although the threats to wild condors persist. Releases began in 1992, and in 2003 the first nestling was reared in the wild. By 2014 the total population of condors was 425 (219 in the wild, 206 in captivity). However, even though condor numbers grew, some individual birds were born with a lethal form of dwarfism. The tiny residual population held only a fraction of what must once have been the condor’s genome, creating a “genetic bottleneck” (when a sharp reduction in the number of individuals reduces the genetic diversity of a population). Despite extensive genome sequencing, it has not been possible to identify a gene or genes responsible for the disease, and a solution to the genetic problem has remained elusive.24



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From the point of view of genetics, evolution can be thought of in terms of changes over time in the relative frequency of inherited traits in a population of animals or plants (or any other kind of organism). Genetic variability within a species allows adaptation to changing environmental conditions, such as the warming of the Northern Hemisphere climate after the last glaciation twelve thousand years ago, or changes in the types of predators lurking in the wildwood or the cerrado. In time, adaptations of a population (species are made up of multiple populations) will be so extensive that a taxonomist would say that that population was sufficiently distinct so as to have become a new species. This has happened when species become isolated, for example, on remote islands, as Charles Darwin famously observed in the Galápagos. Evolution requires variation on which natural selection can act: it is often described as “descent with modification.” Evolution itself reduces variation that is inherited: only those traits persist that confer fitness in an individual or do not negatively affect survival. Individuals and the DNA underlying the traits they possess (longer fins, larger ears, greater density of roots. and so on), are subject to natural selection through mechanisms like freezing winters, dry summers, or competition for food. Natural selection is the process by which individuals with specific advantageous traits leave more offspring than individuals that lack these traits. If those advantageous traits are passed along to offspring— that is, if they are heritable—then that trait will increase in frequency in the population. In species that reproduce asexually, like bacteria (which divide), or many plants (which reproduce through bulbs, tubers, or rhizomes, or by sending out adventitious shoots that take root), all the genes in offspring come from one parent, and all individuals are identical. They are effectively clones. This can give rise to colonies of genetically identical individuals derived from a single ancestor. These can be extremely vulnerable to novel threats such as diseases—British elm trees are clones, and when Dutch elm disease hit the United Kingdom in the 1970s it spread extremely rapidly through woodlands and hedges, killing almost every tree.25 In species that reproduce sexually (as in most eukaryotes, including flowering plants, butterflies, and people), offspring of the same parents are

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not genetically identical (unless they are identical twins, which arise when a single egg splits early in development). This is because every gene comes in two versions (called “alleles”), one from each parent. In some cases, each parent provides the same alleles—in which case the offspring is described as “homozygous” for that gene (from the Latin word homo, meaning “the same”). In other cases, the offspring gets one allele from one parent and another allele from the other parent, making them “heterozygous” (hetero, meaning “different”). Sexual reproduction therefore allows the continual re-sorting of alleles in offspring and the potential to generate more and more variation—and not just any variation but the key to evolution, heritable variation—not suntans or tattoos but disease resistance and height. Changes occur in DNA through a variety of mechanisms, most often through mutation in the process of DNA replication. This replication process is generally very accurate, but not always: sometimes the wrong base pair is inserted. While the nucleus contains enzymes (proteins created from another part of the genome) that detect and correct mistakes, about one in every million nucleotides is copied incorrectly. So in a cell with four billion base pairs of DNA (a corn plant, for example), there will be about four thousand novel pairings, or mutations, every time a cell divides. These are critical generators of variation on which natural selection can act, since the traits they code for may be advantageous or disadvantageous for the individual within which they occur, increasing or decreasing its chance of surviving long enough to reproduce and pass on the new genetic code. This process of novelty creation is happening continuously, in every gene in every cell of every individual of a population of animals or plants, and the process of selection between traits is endless and open-ended. If a mutation happens in an ordinary cell (what biologists call a somatic cell), then it is limited to cells that develop from the original in that individual. If it happens in a gamete, or sex cell, it is what is called a “germline mutation” and is passed on to all offspring. Modification of the germ line obviously has a much greater impact on determining the future genomes of a species. Mutations occur all the time, and most of the time they appear not to make any major difference to the organism. Sometimes mutations greatly increase the chances of death, in which case they do not persist (after all,



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dead animals or plants no longer reproduce!). Occasionally, a mutation occurs and serves to increase an organism’s “fitness” (meaning the success with which an organism gets its genes into the next generation), for example, a plant that is more tolerant of drought or salinity in a desert climate. Because the action of natural selection on genetic variation acts through reproduction, the faster the generations turn over, the faster natural selection acts, and therefore the faster evolution takes place. As a result, microbes with generation times of minutes, hours, and days—the familiar coliform bacterium Escherichia coli divides every twenty minutes—evolve more rapidly than in animals like koalas, which have a generation time of about eight years. Changes in DNA can also be caused by things that damage or break the double-helix structure. The cell attempts to repair damaged DNA, but such repairs are not completely effective. In nature, UV light from the sun and ionizing radiation can do this. Pollution from radiation leaks, for example, in poorly managed nuclear waste dumps, nuclear bomb tests, and the sites of nuclear accidents such as Chernobyl in the Ukraine and Belarus, is a recognized source of DNA damage. The resulting changes can have direct or indirect effects, inducing mutations, causing mistiming of biological processes within the cell processes, and direct damage leading to mutations and sometimes cancers. Scientists have also invented genotoxic chemicals that deliberately damage DNA in target organisms, for the purposes of genetic engineering. Unrepaired DNA may accumulate in nonreplicating cells, such as cells in the brains or muscles of adult mammals (among other things, they are implicated in aging). As mutations accumulate from errors in DNA replication or environmental effects, tissue “mosaics” have been shown to form in the human body. Analysis of RNA-sequence data held within large existing databases has shown a mosaic of clusters of cells with different genomes in many different parts of the human body.26 Genetic variation can also be generated by the action of “epigenetics,” through processes such as methylation (the addition of a CH3, or “methyl” group, to DNA)—not by changes in the genetic code itself but rather by modification of the way genes are expressed. The concept of gene expression can be thought of in terms of the gene being turned “on” and “off” like a switch in an electrical circuit. If a gene is turned “off,” then its

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function is silenced. An expressed gene is active while an unexpressed one is inactive. Though the switching on and off of genes might seem strange, it is common in many organisms. Life begins with undifferentiated cells that must develop into all the parts of the body: the classic blue-and-yellow macaw of the Brazilian cerrado starts as a single cell that must develop into the cells of skin, heart, muscle, gut, and eyes, let alone the cells that form the brilliant yellow feathers of its underside and the vivid blue of its back and wings. This development is accomplished by the expression of some genes in cells and the silencing of others. Scientists have begun to discover that DNA methylation is part of an organism’s response to changing environments, and as such it can affect biodiversity from genetic diversity to ecosystem responses. One study showed that in experimental settings epigenetic diversity actually increased the productivity and stability of plant populations. This emerging field only emphasizes the importance of cross-scale linkages in evolution and the importance of considering them in conservation.27 The “experience” of a gene can also influence its expression. For example, both the nutritional status of parents and stress while a fetus is in the uterus have been shown to change the ways that genes are expressed. Scientists are still debating the significance of epigenetics, but it seems to explain some previously unexplainable facets of evolution. In terms of conservation, there are strong suggestions that epigenetic changes may facilitate the successful invasion of introduced species.28 The range of genetic variation in a population is likely to rise if an individual arrives from a population with a slightly different genetic makeup. Organisms on remote islands, or separated by a long distance from other areas of suitable habitat, tend to have a limited range of genes in their population. The arrival of vagrant individuals that interbreed with residents can introduce new genes. This can be a good thing from a conservation perspective if, for example, the isolated population has started to lose fitness because the gene pool is too small, or a bad thing if the new genes water down the isolated genome and produce a genotype less well adapted to the habitat. Genetic changes are increasingly being understood to take place not only at the level of the organism or the population but at the ecosystem and landscape level. Flying under a variety of names, including community



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genetics, landscape genetics, and landscape genomics, research is exploring the ways in which larger biological and physical patterns affect the makeup and function of genes and the ways these, in turn, influence the ecosystem. Research has shown, for example, that genetic differences in trees can affect the ecosystems in which they are found, including arthropod community structure, aquatic fungal diversity, and ecosystem processes like nutrient cycling. Genetic variation within a species can result in more ecological change than variation between species. And these changes can be very rapid, taking place even over single generations.29 The patterns revealed by genomic analysis at the scale of landscapes have important conservation implications, for example, affecting the way infectious diseases spread and differentially influence host species, how roads impact evolutionary processes, and how landscape modifications affect adaptation in organisms like bees.30 Genes by themselves are a vital part of how life works, but genes do not exist by themselves; they are part of a much larger, linked whole—the genome. Definitions of “genome” vary, but the term basically refers to the complete set of genes or genetic material in a given cell or organism. The word was first used in 1920 by the German botanist Hans Winkler. In the computer-tinged language of modern molecular genetics, it can be thought of as the entire set of instructions necessary to make, run, and reproduce an organism and includes the interactions between the entirety of the genetic material and the environment. In this genomic way of thinking, the inheritance system based on genes is only one of the mechanisms that bring about continuity across generations interacting with epigenetic, cellular, and systematic mechanisms.31 Understanding of genetics has been revolutionized by the rise of genome sequencing in the 1970s and 1980s. Sequencing involves the determination of the sequence of all the nucleotides of all DNA within an individual organism. In the 1970s, DNA sequencing methods were primitive, manual, slow, and expensive, involving the use of chromatography. In the 1980s these became progressively more sophisticated, more automated, faster, and cheaper. By the 1990s, analysis machines and computing power had advanced so far that the sequencing of whole genomes began to be possible.

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The first entire genome sequenced was published in 1995, a bacterium (Haemophilus influenzae, mistakenly thought until the 1930s to be the cause of influenza, which is in fact caused by a virus). This was a relatively small genome (approximately 1.8 million base pairs). The first eukaryotic genome was published in 1996, the yeast Saccharomyces cerevisiae, with about 12 million base pairs. The first multicellular organism to be analyzed was a soil nematode in 1998. The year 2000 saw two staples of molecular genetics sequenced, the plant Arabidopsis thalania and the geneticists’ favorite, the common fruit fly, Drosophila melanogaster. The genome of the laboratory mouse was completed in 2002. It was, however, the sequencing of the human genome that drew public attention to this area of science. One reason is that it was, in effect, a race between a private corporation (Maryland company Celera Genomics, headed by the scientist Craig Venter) and the publicly funded Human Genome Project. Work began in 1990 and the first (still incomplete) human genome sequences were published in draft form in February 2001 by both competing organizations. US president Bill Clinton, speaking in June 26, 2000, described the result as “Without a doubt . . . the most important, most wondrous map ever produced by humankind.” Effective completion came in 2004, with the publication of an almost complete sequence of the entire human genome. By 2018, the genomes of over a million individual humans had been mapped.32 DNA sequencing has become the fundamental technology in many areas of molecular biology and medicine, and it is an essential precursor of the kinds of genetic engineering that are now being undertaken. In medicine, the sequencing of the human genome allows investigation of the genetic basis of susceptibility to disease. Gene sequencing is the basis of DNA profiling in forensic science (typically using the DNA in human hair or saliva, but also extending to wildlife species in cases of poaching and illegal trade). Private genetic testing has become a boom industry, claiming to reveal paternity, remote ancestry (does my family have Viking or Native American ancestry?), and susceptibility to disease. In conservation, the application of genome sequencing has so far been limited by its cost and by limited experience of the use of genomic analyses to address conservation questions. However, that is starting to change. The analysis of genomes is a complicated, machine-heavy practice, more



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associated with clean labs than muddy field boots. Relatively few wild species have had their genome mapped—though this is changing fast as costs fall. Conservation attention has tended to focus on charismatic species like the California condor mentioned above, the orangutans, or the giant panda. To celebrate Canada’s 150th birthday, Canadian scientists sequenced the genome of the beaver. Single-celled animals, the microbes, fungi, bacteria, viruses, and others have been underrepresented, even though they are the most diverse group of organisms on earth.33 Genomic mapping of the wild is proceeding apace. In 2009, the San Diego Zoo began a project (“Genome 10K”), to sequence the genomes of 10,000 wild species (by 2018 they had sequenced or begun work on 277 species). The Earth BioGenome Project was set up in 2018 with the aim of sequencing the genomes of all earth’s eukaryotic species over ten years. Each completed genome of a wild species represents a trove of genetic information available for any purpose biologists can dream up.34 After a long and convoluted path, scientists have finally begun to understand the genetic makeup of life and how it functions. That understanding has brought with it a growing appreciation of the diversity and complexity represented in the cells of the earth’s countless forms of animals, plants, and microbes, and of the extent to which the things about the natural world that humans both appreciate and need—the migrating herds of wildebeest and pollination by millions of insects—rely on the complexity of the genes and their ability to respond to changing conditions. Life on earth, including human life, relies on the dynamism of DNA. But rapid global environmental change is placing new selective pressures on species, which must either adapt or migrate to find environmental conditions where they can survive, or they will most likely go extinct. Conservation scientists are increasingly recognizing the importance of evolutionary perspectives in conservation management programs and exploring what Eizaguirre and Baltazar-Soares term “evolutionary conservation,” enhancing the capacity of species to adapt to environmental change. Work to understand DNA and genes is showing its relevance at larger and larger scales and is emerging as of vital importance in conservation practice as we describe in the next chapter.35

4 Conserving the Genetic Pieces

“Look, he’s up there, in the big tree, just behind the Spanish moss.” We craned our heads trying to follow the directions from our guide at White Oak Plantation. We were fifty kilometers north of Jacksonville, North Florida, deep in the heart of what used to be first a rice and then a tree plantation and is now a center for wildlife conservation and endangered species management. White Oak had experience with captive breeding of endangered rhinos, cheetahs, and okapi, when it was asked to work with the highly endangered Florida panther. And that was what we were watching, in a big enclosure deep in the North Florida shade: ten meters off the ground, astride a large live-oak limb, a big panther stretched out. The Florida panther (Puma concolor coryi) is a recognized subspecies of the puma (also called the mountain lion or panther). It was once found across a broad swath of the southeastern United States, yet as early as 1870 it was rarely to be encountered over much of this range, a victim of habitat loss and targeted hunting. It was so rare that some said it was a mythical beast. By the early 1990s there were only a very few left, some twenty to thirty animals, deep in the Big Cypress region of South Florida, but they were not in good shape. Not only were there so few individuals, they had some of the lowest levels of genetic variation observed in wild

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felids and were suffering from problems ranging from poor sperm quality to compromised immune systems and cryptorchidism (absence of one or both testes in the scrotum).1 In 1967 the Florida panther was included on the inaugural US list of endangered species. It was soon clear that conventional conservation strategies (for example, the creation of protected areas or controls on hunting) were not going to save it: the population was collapsing in on itself. In 1992, White Oak Plantation hosted a meeting at which the bold decision was made to try to save Florida panthers by addressing the bottleneck in their genetics. It was recognized that this could not be done by breeding the remaining wild panthers, since they were already too inbred. Genetic help was needed from outside—in this case from eight female panthers from Texas, which were introduced into the Florida wilds in 1995. They bred successfully, leaving some of their genes in their offspring. The Texas females were then removed from the wild, having performed a mission of “genetic rescue.” Assessments of the impact of this remarkable intervention show a strongly positive conservation impact. The wild population has grown, its genetic diversity has increased, and fewer medical problems are reported. It can be said that the Florida panther has been “rescued,” in a genetic sense. Yet even now, further introductions of pumas from other populations (experts suggest releasing five to ten individuals from other puma populations every twenty to forty years) are needed to make the Florida population secure.2 But how should we think about the rescued Florida panther? Pumas, the species of which it is a part, are the most widespread mammal in the Americas, found from Patagonia in the south to Alaska in the north. The history of their classification is long and complicated, with Linnaeus himself providing the genus name—Puma. Only one species is recognized, Puma concolor, literally, “the Puma of one color”—distinguishing it from cats with spots, like jaguars. This seems simple enough, but looking at variation within the species it gets more complicated. In a 1946 publication, the species (Puma concolor) was divided into thirty-two subspecies—one of which was the Florida puma, Puma concolor coryi, and another its genetic savior, the Texas puma, Puma concolor stanleyana. But a 2000 study based on a genetic

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analysis collapsed these thirty-two subspecies into only six, and a 2017 study using an analysis of mitochondria brought this down to only two subspecies, one in South America and the other in Central and North America. These genetic analyses supplanted the previous taxonomy based on measurements of shape and size—particularly of the skull. So where does that leave the Florida panther and the efforts made to conserve it?3 Everyone agrees on what a puma is, but there is evident disagreement on how to classify the variations within the species. And everyone agrees that the injection of genes from Texas pumas have enabled the last wild pumas in Florida to survive. But was this a mixing of different subspecies of pumas (meaning that the “Florida panther” is now a hybrid)? Or was it just a mixing of different populations of the same subspecies? And does this distinction matter, and if so, to whom? Humans like to classify things. We routinely recognize and enumerate differences of all kinds among ourselves and in the world around us: he is a Republican, she is a bird-watcher, that is a geranium, this is Gruyère cheese. In her book Purity and Danger, the anthropologist Mary Douglas argues that classification is so important for the functioning of human societies that it is buttressed by social norms that reinforce it.4 The story of the Florida panther highlights several things about the importance of taxonomic classification in conservation. First, it reminds us of the power of the species (or, as in this case, the subspecies) as a unit in conservation management. Species are conventionally thought of as natural units, sets of discrete nonoverlapping categories. The concept of species has a remarkably long history, extending back to Plato and Aristotle and their attempts to create universal classification systems. Students of biology learn that species are the units of evolution, although if they paid attention, they would remember that it was actually the variation between individuals within the species (or populations) on which evolution acts. Species are conventionally said not to crossbreed—or rather, if two species do, their offspring will not be fertile, as the owners of mules, the infertile progeny of donkey and horse, have known for millennia. Although the need for other ways of prioritizing action is widely recognized, species, whether giant pandas, great white sharks, or bristlecone pines, continue to be the focus of much conservation attention.5



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Second, conservationists commonly assume that taxonomic distinctions are clearly and naturally defined, yet scientists don’t share that conviction, as the disagreement on how to classify puma subspecies shows. There is considerable scientific disagreement on how to classify species. It has always been the case that naming a species is a professional accomplishment for taxonomists and often tinged with scientific politics. People liked having things named after them, and countries liked having species not shared with another country. The result, as Brooks and Helgen point out, has been a trend “towards recognition of progressively smaller levels of differentiation as evidence of species status. Just as monetary inflation can devalue a currency, this taxonomic inflation can erode the usefulness of the species as a robust measure of the diversity of life.”6 The process of naming species has not been a constant one. Changing practices across centuries have often resulted in a “dog’s breakfast” of taxonomy, which is one reason for contemporary restructuring. Applying a consistent taxonomic approach to naming species can bring dramatic changes to the species recognized and, hence, to conservation programs based on them. For example, a comprehensive revision of the taxonomy of all species of birds led to a 10.7 percent increase in the number of bird species recognized, an increase of over one thousand species. The ability to map the genomes of wild species is a major driver of changing taxonomic classifications. Genetic analysis has, for example, split the previously single species of giraffe into three species: the northern species, the southern species, and the Masai giraffe. Such tools have also allowed increased examination of the genetic structure of individuals, populations, and species.7 Third, conservation organizations, like government agencies and international treaty bodies, build legislation and laws around particular species, and their work tends to reinforce a given taxonomy. Lists of rare or endangered species have existed since the 1940s, and the IUCN Red List (founded in 1964) remains the key information source for measuring the risk of extinction and therefore the importance of different conservation actions. Species at risk of extinction often receive extensive conservation investment, both in their own right and as “flagships” for wider groups of organisms and ecosystems. Conservation fundraising depends on the power of widely recognized species (particularly mammals) to motivate public support for conservation.8

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In the United States, the Endangered Species Act (1973), in which the Florida panther is listed, mandates a species-focused view of nature— though as “species” it also considers subspecies, varieties, and, for vertebrates, distinct population segments. It requires all federal agencies to ensure that any action they fund, authorize, or carry out will not jeopardize the survival of any endangered species or degrade designated areas of critical habitat for such species. The law also prohibits hunting or trade of any endangered wildlife or fish species. In the United Kingdom, the model of “species action plans” provided a powerful bureaucratic framework for addressing declines in key endangered species in the 1990s, as the government worked out how to respond to the Convention on Biological Diversity, which it had just signed in 1992. Endangered species provide a powerful structuring device for broad international legislation and cooperation: 183 governments or regional economic integration organizations currently belong to the Convention on International Trade in Endangered Species (CITES), which makes rules on trade in products from wild species, such as elephant ivory, shark fins, and live orchids.9 The species is a human-designated category, the result of careful scientific analysis and liable to change as new scientific findings become available. It is therefore a tricky unit on which to base conservation action. Yet it is also long established, widely recognized, and effective. Above all, it gives conservation a relatively simple handle on the diversity that evolution has created. One example of this is conservation’s particular concern for species that are the only surviving members of now-extinct lineages. These represent an extraordinary library of diversity and a record of the evolution that has been going on since that lonely Last Universal Common Ancestor four billion years ago. In terms of Darwin’s tree of life, these are the last twigs of almost-lost but formerly extended branches. One such species is the tuatara, an unprepossessing greenish-brown lizard weighing about the same as a chicken and found only on a few isolated islands in New Zealand. The tuatara looks a bit like a miniature dinosaur, but it is in fact an unspecialized reptile, with some characteristics of the amphibian line from which it developed more than two hundred million years ago. It has been protected since 1895 in New Zealand, but like so many other endemic species, it suffered greatly from habitat loss and predation by introduced species like the Polynesian rat. By the



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end of the twentieth century it was extinct on the New Zealand mainland, restricted to a series of small offshore islands until the Department of Conservation hatched a plan to reintroduce tuatara to a fenced sanctuary on North Island, from which all predators had been removed.10 There is a genetic logic to the protection of the tuatara that is quite separate from all the cultural reasons that might exist to put great efforts into its conservation. Its genome is quite different from that of other reptiles. As the only survivor of a taxon that split away from others so long ago, it holds a great deal of unique accumulated evolutionary history. This gives it an importance denied to more diverse taxa, such as the fruit flies of the genus Drosophila, the workhorses (in an odd turn of phrase) of laboratory genetics and the denizens of the fruit bowl after your week at the beach. Small fruit flies, also called vinegar flies and pomace flies, belong to a family of flies with almost five hundred genera worldwide, and around fifteen hundred species in the single genus Drosophila. So, unlike the case with the tuatara, one or two species of Drosophila do not contain much accumulated evolutionary history. Unlike the tuatara, the genetic history stored in any single species of fruit fly is of lesser conservation value.11 Conservationists expend a lot of effort trying to protect the integrity of species. Where species intermingle by hybridization, conservationists worry that the natural genetic diversity of distinct species will be lost, and potentially the species itself. Where wild species populations are reduced to small numbers, as happened to Florida panthers, conservationists worry that the capacity of that species to have the genetic flexibility to survive will be impaired. With captive populations, animals are moved between zoos to keep healthy gene pools and prevent inbreeding. Yet there is still no agreement on how one species is to be separated from another; a recent compilation lists thirty-two different ways of defining a species. And this doesn’t even include the difficulties of applying the species concept to microbes, which leads some to question the very utility of the concept. In fact, this conclusion may apply to the category of species itself, with the author of one review ending his paper by saying, “perhaps the most fundamental issue is whether the term ‘species’ refers to a real category in nature.”12 The recent hubbub around our own species, Homo sapiens, is illustra-

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tive. Powerful genetic technologies, able to recover DNA from fossilized bones, have revealed that humans interbred with at least two other species, Denisovans and Neanderthals, and perhaps others yet to be unmasked. In fact, one fossil was found whose DNA revealed a tryst between a Neanderthal mother and a Denisovan father. This pattern of breeding between hominid species is being revealed to be much more common than thought, and the notion of humans as a unique, inviolate category continues to spring leaks.13 The same complexity is true in other species. As more genomes are deciphered, more and more hybrids are being revealed. Genomes are increasingly understood to have porous barriers. In animals, sharing of genetic material across taxa has been shown to occur both in the present and in the evolutionary past—found in creatures ranging from fish, wolves, and frogs to corals, elephants, and crocodiles. Plants take the prize, however, with up to a quarter of plant species thought to be of hybrid origin. Species of forest trees often hybridize across zones where two species meet, a phenomenon found in taxa as diverse as eucalypts, poplars, oaks, and spruces.14 Hybridization is not the only way genes can slip between apparently distinct species. Individuals in a species are conventionally thought to receive their genes from their parents, who in turn received them from their parents, and so on back to that first common ancestor. However, that is not the only way genes can end up in an organism’s genome. Genes can also sneak in through the back door of cells, for example, when genetic material is passed directly from one genome to another without the usual pattern of parent-child inheritance. Labeled “horizontal gene transfer,” or a related term, “gene introgression,” this mechanism appears to be much more common than biologists thought until recently, particularly in bacteria. In bacteria, swapping genes appears to be a reasonably common way of life, and horizontal gene transfer is a major driver of their evolution. The mechanism has allowed bacteria to rapidly develop resistance to powerful antibiotics as well as to grow in a wide range of environments—including cheese rinds. A recent study revealed that 80 percent of the 165 bacterial species found on cheese rinds shared genes with other species. Nearly five thousand genes had been exchanged.15 Technical advances in the reading of genomes have led to a growing



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realization that bacteria have also been important sources of new genetic sequences for multicellular animals and plants. Almost one in twenty flowering plants appear to carry bacterial DNA in their genomes, including sweet potatoes, American cranberries, and even the hops used to make beer. Genes are also shared between related species. In mushrooms, the genes coding for psilocybin, the psychoactive compound sought by hippies in the 1970s, were shared between at least eleven distantly related mushrooms through horizontal gene transfer. Recent work on grass genomes has shown that large blocks of DNA containing functional genes have been passed horizontally between distantly related species.16 It is not just plants whose genomes have been enriched by bacterial genes; recent work published in 2015 suggests that humans have up to 145 genes in their genomes that jumped from bacteria, other single-celled organisms, and viruses. However, most genes that originated outside the human species are nonfunctional and end up in what one author has referred to as a “junkyard” of other genes.17 But one organism’s junkyard may be another organism’s treasure. Recent work has shown that brand-new genes, de novo genes—never known before and unrelated to genes in similar organisms—can arise with considerable frequency, seemingly from noncoding parts of the genome. Such “junk DNA” has been found in many species ranging from fruit flies or fish to crop plants, as well as to humans.18 And one more trick biology has developed to move genes around in not-your-mother’s-usual way is through something called “jumping genes,” or, more formally, “transposable elements” or “transposons.” First identified by geneticist Barbara McClintock, transposons are DNA sequences that have the ability to change their positions in the genome. There are many different types of transposons, and they make up the great majority of a genome’s junk DNA. They can occupy a significant portion of the genome: in corn, for example, transposons make up 60–70 percent of the maize genome.19 Not only can transposons move within genomes, they can move across genomes. Such movements have been detected in many species. Most have been identified in animals, but transposons have also been shown to have crossed the species barrier from host to parasitic plants. They can also move between very different kinds of species, for example, between

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an arthropod and a plant. In another remarkable cross-species jump, the transposon BovB—which, as might be guessed from its name, was first discovered in cows—has been shown to have moved to reptiles, amphibians, bats, elephants, and marsupials, with ticks the probable agent facilitating this movement. So the genome is not a calm, quiet place where nothing changes, but is a churning mass of genetic material, occasionally—maybe more often than just occasionally—moving from one type of organism to another. But it is not just the diversity of methods of gene exchange that undermines conventional ideas about discrete genomes belonging to discrete individuals belonging to discrete species. There are also genetic exchanges between host organisms and the species in their microbiome, the species living within and enrobing the bodies of animals or plants. These interact with the host in powerful ways across species boundaries, comprising a sort of hyper-genome. Some of those interactions are physiological (through the ecosystems of gut, root, or skin). Some are genetic. The composition of the microbiome changes in response to hygiene, medication, diet, and proximity of organisms of the same and other species. The microbiome is also constantly changing due to exchange of genetic material between microbes. Bacteria that form part of human microbiomes have a twenty-five-fold higher rate of gene transfer than do bacteria found in other settings. The complexity and range of these interactions are such that geneticists Elizabeth Grice and Julia Segre have suggested that the microbiome should be considered a “second genome”—not entirely separate but interacting in many ways across species boundaries.20 Microbe communities are not genetically independent of their host species or of the ecosystems within which they mutually exist. Recent work in Hawaii has shown complicated interspecies interactions in a single valley, in what the microbial ecologist Anthony Amend calls an “ecosystem microbiome.” The microbiome of each species of animal and plant in the valley consists of a subset of what exists in the whole ecosystem: no microbiome is an island but a part of a whole. Such connections between microbiomes extend between realms, with recent work showing movement between airborne microbes above the Great Barrier Reef and those found in the microbiomes of corals beneath the ocean.21



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The variety of life so valued by conservationists exists not only in the genes and genomes of organisms, but also in the genes of microbiomes. As we will describe in Chapter 6, introgression between species, and their microbiomes, is enormously important as scientists start to engineer genomes. It becomes harder and harder to isolate where one organism’s genome begins and ends. Genes, species, and ecosystems are the result of the dynamic forces of geology, climate, evolution, natural selection, and extinction. They are intricately linked to one another in ways we are still learning about. Conservationists are dedicated to protecting and maintaining this diversity. But genomes, like ecosystems, keep changing. When students are taught evolution, they are usually told to imagine the process taking tens of millions of years, like the eventual evolution of whales from a four-legged goat-sized amphibious animal called Pakicetus. Evolution as conceived by Darwin was something that was long and slow. It turns out this is true but not sufficient: evolution can also take place rapidly and is doing so now all over the earth. Humans have come to play a growing role in shaping its course. In the process of converting the earth to meet their needs, humanity has become a major driver of evolutionary change.22 Moreover, human activities, among them pollution, climate change, agriculture, hunting, fishing, and drug development, all influence the genetics of wild species. Human action has also increased the incidence of hybridization between species because the conditions created by humans are different from those under which the species evolved. For example, the creation of novel ecosystems (including novel microbiomes) creates conditions wherein traditional reproductive boundaries can break down. Novel genomes and, with them, novel microbiomes are becoming part and parcel of novel ecosystems.23 The effects of harvesting on the genetics of wild animals and plants have been particularly well studied. Relentless commercial fishing, for example, decreases body size in fished species by over 300 percent and decreased the age at which these fish reproduce. In other words, catching big fish results in genetic changes in those that remain, making them smaller and causing them to breed earlier. Even the milder pursuit of rec-

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reational fishing, dropping a line in a nearby brook or pond, can change the morphology of fish, resulting in fish with smaller mouths—less able to ingest the dangerous hook.24 In other cases, species have evolved to cope with living alongside humans. In northern Europe, for example, open heathland is spoken of as a domesticated ecosystem. The habitat is created by human land-use decisions over thousands of years. This is the classic open landscape of Jutland in Denmark, maintained by cutting (of pine and birch trees and other plants), burning, and grazing. It is dominated by common heather, which is good at regenerating after fire. Recently, researchers showed that heather from traditionally burned coastal heathlands in Scandinavia had an adaptation that meant that it was stimulated to germinate by chemicals in smoke, while heathlands with infrequent natural fires did not. Human influence seems to have shaped not only the heathland ecosystem, but also the genetics of its dominant plant. Heather has a partially domesticated genome.25 Wild species have also undergone genetic changes to take advantage of the environmental modifications wrought by humans. Evolutionary biologist Mark Ravinet and colleagues looked at the house sparrow, that almost ubiquitous suburban and urban bird. House sparrows feed on food waste and crops. They are fully dependent on human-caused changes and probably have been since the emergence of agriculture in the Neolithic Revolution. Researchers compared the genomes of three Eurasian sparrow species that are all dependent on humans (the Spanish, Italian, and house sparrows) with the bactrianus subspecies from the Middle East and Central Asian steppes, which is not associated with human settlements and which migrates. Modeling suggested that the house and bactrianus sparrows diverged about eleven thousand years ago, and the house sparrow expanded its range and population size about six thousand years ago, at the time of the expansion of agriculture into Europe. A comparative genome scan identified genes potentially involved with adaptation to eat a higher starch diet as well as to have a more robust skull that allowed a shift from wild food to specialize on the resources provided by agricultural crops. On the strength of these genetic changes, house sparrows have spread to almost every continent on earth.26 A more contemporary setting—the city—is also playing a part in driv-



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ing rapid evolution of wild animals and plants. More than half of the earth’s human population lives in cities, and urban theorists talk about a  process of “global urbanization,” as urban economies and associated technologies and lifestyles extend their tentacles of production and consumption into even remote regions. Urban environments are recognized by biologists as hotbeds of rapid evolutionary change. In the words of Menno Schilthuizen’s highly successful book Darwin Comes to Town, the “urban jungle” is now “driving evolution.”27 Artificial lighting, higher temperatures, more cement and less soil, changed water flows, and reduced air and water quality from pollution all change the environments to which wild plants and animals adapt. It is well known that urban areas provide novel habitats containing non-native species that compete with native species for space and resources. However, urbanization can also affect the evolution of species by increasing the rate at which genes mutate, by changing gene flows through isolating populations (think of a population of white-footed mice isolated in a city park), and by changing the evolutionary pressures on wild species in unexpected ways.28 Even something as commonplace as a streetlight is not as evolutionarily innocent as it might seem. The evolution of nocturnality—being active in the dark—is found across many kinds of animals and is based on the constancy of a roughly twenty-four-hour cycle between darkness and light. So when streetlights, stadium lights, airport lights, or even porch lights break this never-before-broken cycle, it can change a lot. Recent work by Kevin Gaston of the University of Exeter and his colleagues has shown how disruptive night light can be to nocturnal creatures and how it is affecting their evolution. Artificial light at night can influence the evolution of body size, immune function, and photosynthetic rates. Light pollution is also a problem for the seaside, where light pollution has been shown to significantly stress corals.29 Another example of rapid evolution to human-induced selective pressure comes from the widespread use of chemical pesticides. The evolution of pesticide resistance in weeds, pests, and parasites is becoming a huge economic problem (as anyone with a child in a class where someone has lice can attest). Human-made poisons put huge evolutionary pressure on target species so that any mutation that confers an enhanced chance of

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survival is likely to be selected for strongly. This drives the evolution of resistance to the poison, and an expensive and endless arms race by the pest-control industry to develop novel and more toxic poisons. The best-documented case of human activity driving evolution comes in the rise of microbial resistance to antibiotics in human medicine. In this case the human body is the ecosystem, and bacteria are the wild species. The use of antibiotics to control infections in the human body is a standard tool of medicine. Yet in the United States alone, at least two million people are infected with antibiotic-resistant bacteria and over twenty thousand die as a result. Antibiotics are heavily used: in the United States in 2015 medical professionals wrote 270 million antibiotic prescriptions. Research suggests that a significant number of prescriptions are inappropriate. But in animal agriculture the use is even higher. In 2016, in the United States alone, approximately 8.4 million kilograms of antibiotics regularly used in treating humans were used in livestock production.30 The problem of antibiotic resistance is well known. It is less often framed as a case of extremely rapid evolution. In one infected patient being treated with the antibiotic vancomycin, doctors documented bacteria undergoing thirty-five separate mutations to become resistant. Under intense selection from antibiotics, the movement of genes from one bacterial species to another without reproduction (horizontal gene transfer) speeds up the rate at which resistance evolves. The human body provides a neat, if scary, model of how extensively evolution can be shaped by human action.31 It is clear that the genomes of wild species are not static but changing, both in the short term and long term. Moreover, humans have become major sources of such change. Many of the changes humans cause are inadvertent, but an increasing number are deliberate. An increasing number are being made by conservationists themselves, as they develop techniques to manage the genomes of species they are trying to protect. Decades ago, it became clear that one technique to preserve the genetic diversity of rare species was to remove them from the factors that threatened their extinction. Termed ex-situ conservation, this involved removing individual animals, plants, seeds, or cuttings from the ecosystems where they occurred, either to store them or to reproduce them in



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controlled conditions. For plants, the main approach has been to create seed banks, drying and storing seeds under conditions of low temperature and moisture. These have been widely used in plant breeding since the 1930s, to preserve the wild relatives of domesticated plants or old “landrace” varieties of historical and cultural value for future breeding endeavors. Now there are much larger public facilities to preserve the seeds of wild plants, notably the Svalbard Global Seed Vault, drilled deep into the rock in remote Svalbard, Norway (an Arctic territory), and the Millennium Seed Bank, operated by Kew Gardens in southern England, which aims to obtain and preserve all the earth’s plant species. A similar approach is now used for wild animals. Since they have no seeds, samples of genetic material (tissue, sperm, eggs, embryos, and DNA) are frozen in tanks of liquid nitrogen. The San Diego Zoo in the United States has set up what it calls a “Frozen Zoo” (a term they have copyrighted), which holds living cell cultures, oocytes, sperm, and embryos from almost one thousand taxa. They even have genetic material from an extinct species, the po‘ouli, or black-faced honeycreeper, a bird formerly endemic to the island of Maui in Hawaii. The Frozen Zoo is intended to serve as a resource for research and for conservation, for example, in assisted reproduction (such as in vitro fertilization, artificial insemination, and embryo transfer). However useful and ingenious, gene banks can only preserve individual species—and really only their genotypes, which are only “parts” of the overall species and don’t conserve their interactions with other species and the ecosystem, or their microbiomes. They fix nature’s diversity at a particular point in time, imposing an “evolutionary freeze.” Outside the gene bank, the living world from which genetic material was taken goes on evolving (pests, weeds, pathogens, and all), but the protected genome does not.32 Ex-situ conservation also includes populations of animals and plants kept in botanic gardens or zoos. In accredited zoos with conservation programs, such populations have long been managed using genetic criteria. But since the number of animals of a species kept in any zoo is relatively small, the key to maintaining a viable nucleus for reintroduction depends on three things. First, all animals of a single species need to be managed in an integrated “captive population”; second, careful records

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need to be kept on how different animals are related to each other (since most zoo specimens come from other zoos); and third, animals need to be moved between zoos to breed to maximize genetic diversity. Traditional techniques have allowed zoos to take pioneering roles in restoring wild species, including the American bison, California condor, black-footed ferret, and Panamanian golden frog. The Arabian oryx is a prime example of this process in action. By 1962 this species, found in the deserts of the Middle East, was effectively extinct in the wild. There were two small herds of oryx in captivity, one in the Phoenix Zoo in the United States and one in Riyadh in Saudi Arabia. Captive breeding proved effective, and a “World Herd” was declared, with a “stud book” managed by an international consortium. Currently there are over six thousand captive Arabian oryx worldwide. Several attempts had been made to reintroduce the species to the wild in the Middle East. One of the most significant problems that conservationists have worked to address is the loss of genetic variation in wild populations. The longterm survival of a species depends on the genetic variation found across the many individuals within it. Individuals with slightly differing genetic composition can respond differentially to stresses, and those able to cope produce offspring better able to thrive. When the size of a population drops significantly, the species loses much of its genetic variation—and therefore its chances of survival. The much-diminished populations of the Tasmanian devil have low genetic diversity and face increased susceptibility to disease. The even-more reduced populations of Iberian lynx in southwest Europe show significantly lowered reproductive ability due to poor sperm quality and compromised immune systems.33 Conservation geneticists have been aware of the problem of reduced genetic diversity for decades, particularly those charged with the survival of small populations maintained in zoos. Techniques have been developed to maintain or increase the genetic diversity of isolated populations in the wild, developing the approaches used in zoo collections to maintain genetic diversity. These approaches are often termed “evolutionary rescue” or “genetic rescue.” The story of the Florida panther, at the start of this chapter, is an example of this approach. There are many other examples: pregnant bighorn sheep ewes have been moved to another population of sheep to increase the genetic diver-



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sity, and European vipers in an isolated population in Sweden were “rescued” genetically when vipers from a healthy population were introduced. This approach may also involve applying contraception to certain individuals in captivity, in large enclosures or even in the wild, whose genetics are overrepresented in the population. This has been proposed for several species of marsupial in Australia including black-flanked rock wallabies, and burrowing bettongs.34 But moving genes doesn’t always mean moving animals and waiting for reproduction. It can also be done by bringing females into captivity, mating them with captive males, or impregnating them with the sperm of genetically desirable males and returning them to their wild homes. Instead of moving the animals, their eggs, sperm, or pollen can be relocated in order to increase genetic diversity in a population with low genetic diversity. Any of these techniques of genetic rescue can potentially achieve the desired outcome of increasing genetic diversity.35 In California, salmon have been so reduced in number and hatchery practices have created so many inbred fish that genetic rescue has been customized in an extraordinarily intensive fashion. When individual fish return to the hatchery they are captured and have a piece taken from a fin; then that is FedEx-ed to John Carlos Garza at the University of California Santa Cruz, who analyzes each salmon’s genetics and proposes, in what he calls a “salmon mating service,” which pairs are genetically least related and should breed.36 Genetic rescue has now been practiced for plants, invertebrates, fish, birds, reptiles, and mammals. However, genetic rescue is not risk-free. Some people are concerned about the loss of species identity through hybridization. So, with the Florida panther, there are wild animals abroad in Florida, but with a genome blended with Texan animals. At the same time, hybridization preserves part of the original genome that would otherwise have been lost when a population of species is at risk of extinction (as in the Florida panther case).37 Hybridization also occurs naturally. It has been shown to have happened in bears: part of the genome of the now-extinct Pleistocene cave bear lives on in grizzly bears. But in many cases, hybridization, the mixing of two distinct yet closely related taxa, is seen as a threat to endangered species, resulting in “extinction through hybridization” or “extinction

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through introgression”—the collapse of species boundaries as the result of the transfer of genetic information between species.38 Hybridization is, in fact, found in a wide range of species. It is widely recognized to be a particular conservation problem in the dog family (Canidae). One example is the wolf. Over thirty closely related subspecies are recognized, from the New World Arctic to India (as readers of The Jungle Book will well know). The wolf went extinct in much of its range, particularly in Europe, although it is coming back fast—and meeting domestic dogs almost everywhere. Domestic dogs share over 99.5 percent of their genome with wild wolves—indeed, the domestic dog is classified as a subspecies of the wolf (subspecies familiaris).39 It is no surprise to find that crossbreeding is common. It is not just Jack London’s White Fang that had a wolf as sire: many a sled dog has wolf genes, and the interbreeding of wolves with domestic dogs is widely reported across Europe. In the Apennines of Italy, dog and wolf genes are inextricably mixed in the small population of apparently “wild” wolves, and wolf × dog hybrids have become a serious conservation issue. In the eastern United States, hybrids of coyotes and wolves occur widely.40 As with dogs, so with cats. Domestic cats and European wildcats are classified as separate species within the family Felidae, but they interbreed freely and produce fertile offspring. Wildcats have a very wide distribution, with subspecies across Eurasia and North Africa. In the United Kingdom, they are now restricted to remote areas of Scotland, where the “Scottish wildcat,” after centuries of persecution as a predator, is now an important target for conservation action. Domestic cats are thought to have evolved from wildcats some four thousand years ago, and genetic analysis suggests that as they spread across human societies they continued to interbreed with local wildcat populations. In Scotland, they still do so, and the dilution of wildcat genes by crossbreeding is regarded as a critical issue in efforts to maintain the species. Many large tabby cats living wild are held to be wildcat × domestic cat crosses, and there are detailed guidelines for trying to identify critical features of striping and tail that allow them to be distinguished. Such features do not necessarily have a clear genetic basis, with the result that the distinction drawn between wild and domesticated is arbitrary, even if regarded as of great importance.41



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Human alteration of ecosystems through such activities as construction of roads, changing drainage patterns, and coral removal has resulted in hybridization in taxa ranging from plants and fish to chipmunks and frogs. A recent example involved the adaptation of the Gulf killifish to survive in the highly polluted waters of the Houston (Texas) shipping zone. This species has evolved rapid resistance to the pollutants based on the introgression of a gene from a closely related species, the Atlantic killifish. In a terrestrial example, scientists have documented the rapid spread of rodenticide (warfarin) resistance in a subspecies of house mice based on a single gene that introgressed from the related Algerian mouse.42 Despite hybridization’s acknowledged downsides, there is the beginning of a movement to value hybridization in some circumstances as the opportunity to increase diversity and ability to adapt to a changing world. After all, hybridization can be an important source of genetic variation on which natural selection can act. Deliberate hybridization, or as it is more officially called, “adaptive introgression,” has started to be talked about as a conservation strategy.43 Deliberate adaptive introgression may be a particularly important tool in rapidly changing environments where existing genetic variation and mutation rates may not be sufficient to allow for adaptation. This includes places with novel and rapidly spreading diseases. Paul Woodcock and his colleagues have suggested that trees or their seeds that have a genomebased resistance to disease could be moved into a region where disease is affecting local populations. This has been proposed with ash trees, threatened by ash dieback disease in Europe, caused by a fungus introduced from East Asia. Hybrid trees from zones of natural hybridization might also be moved in anticipation of climate change to increase adaptive genetic diversity.44 It has also been suggested that genetic introgression might be used as an adaptive response to anthropogenic climate change. To this end, researchers have created a hybrid of two coral species from Australia’s Great Barrier Reef in the laboratory. Unlike the two pure coral parents, some individuals in the early generations of the hybrid showed higher survival rates and normal physiological function in the presence of warmer waters and elevated CO2 levels.45 Work such as this on trees, corals, and many other species is only pos­

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sible due to the rapid growth in technologies for reading DNA and mapping genomes. These are transforming the understanding of the biology of species and their genomes and beginning to make possible previously unheard-of types of management. Among other developments, these technologies are allowing species to be identified by their DNA alone, without other physical evidence of their presence in an ecosystem—knowing without seeing. Sequencing allows the identification of short lengths of DNA that are unique to a species. These are known as “barcodes,” and reference libraries allow rapid species identification from DNA alone. DNA can be taken from biological samples (hair, for example), just as it is in human forensic science. It is becoming commonly used to distinguish endangered species in the wildlife trade. Traders go to great lengths to hide the identity of illegally traded species, for example, moving eggs instead of adults, shipping seeds instead of adult plants, or claiming that elephant ivory is from mammoths. In all these cases the claims can be tested using such DNA barcodes.46 Species can also be identified from environmental samples of water, snow, soil, or even air. Such “environmental DNA” or “eDNA” is defined as genetic material that is obtained directly from environmental samples (for example, soil, sediment, or water) without any obvious signs of biological source material. It provides a powerful alternative to traditional monitoring techniques, particularly in the identification of cryptic species or juvenile life stages or invasive species. eDNA can be used to detect single species or to analyze the composition of ecosystems. It can be extracted from sediments (for example, to identify extinct species) as well as contemporary samples.47 New tools allowing the analysis of genes and genomes also make possible the deliberate curation of genomes, for example, to minimize inbreeding and loss of genetic diversity, manage wild populations, and establish genetic-based breeding programs for endangered species. Conservation science journals in the 2010s have been thick with papers using genetic analysis to reveal different aspects of the biology of rare species, straining to identify some relevance of their discoveries to conservation. The science may be outrunning conservation management. But that is changing rapidly. In 2018 Rebecca Johnson of the Australian Museum



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Research Institute and colleagues published the genome of the koala. Their work revealed the genetic basis for koalas’ ability to detoxify eucalyptus leaves, and it identified separate populations with low genetic diversity, highlighting the need for habitat corridors to maintain regional gene flow. In another example, analysis of the genome of wild bees is contributing to conservation and management by identifying parts of the genome associated with disease susceptibility, responses to environmental stressors, and the diversity of associated beneficial microbes.48 An extraordinary example of the application of knowledge of the genetics of a wild species to its conservation is the northern quoll of Australia, a small carnivorous marsupial. These fierce carnivores are highly susceptible to the poison carried by invasive cane toads, and quolls’ numbers have been decimated by their unfortunate habit of trying to eat the toads as the invaders advance. However, there are individuals in these populations that appear to have a genetically based behavior that means they do not bite into the deadly toads. It has been suggested that these toad-smart quolls could be used to spread the genes for that restraining behavior and thus reduce their population losses, or even that these animals’ genes associated with toad-avoidance could be implanted in virgin quolls.49 So, people are changing the genomes of wild species in many ways, not all of them obvious. Genomes, both of species themselves and of their microbiomes, just like ecosystems, are becoming “novel.” Against a background of rapid evolution and continual change in genomes, conservationists are becoming increasingly creative in their struggle to maintain the diversity and integrity of the genomes of wild species, using insights from advances in genomic science. Some conservation practitioners are deliberately mixing genomes in the name of saving species, as they did with the Florida panther. In the process they are knowingly crossing the boundaries between species or subspecies that have so long been the foundation for much of conservation. As Hamilton and Miller say, “to maximize adaptive potential in response to changing environmental conditions, purposeful propagation of genetic variation via human-mediated hybridization may be necessary to conserve at-risk species.” They are not alone in this position. Chan and co-authors conclude a recent paper, saying “perceptions of potential risk

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change significantly when the focus of conservation is on preserving the adaptive potential of a species/population, instead of preserving the species in its original state.”50 All the problems of naturalness, artificiality, and novelty that are familiar in the context of ecosystems also extend to the genetic level. For conservationists, this is a challenge, because they mostly have less training in thinking about genomes than they do about ecosystems. The public that supports conservation also tends not to know much about the genetic dimensions of diversity. Yet the genome—and its diversity, changeability, and openness to deliberate management—is taking an increasingly important position in conservation thinking. Now, new scientific advances, including the capacity to manipulate the genomes of living organisms, are about to make debates about naturalness at the genomic level a whole lot more urgent—and much more complicated.

5 Rewiring Nature

On May 21, 2010, Craig Venter, the scientist known for mapping the human genome, stood on a platform at a museum in Washington, DC. In a calm voice, he announced his team’s creation of a “synthetic cell” or “the first self-replicating species we have had on the planet whose parent is a computer.” This involved, he said, “starting with the digital code in the computer, building the chromosome from four bottles of chemicals, assembling that chromosome in yeasts, transplanting it into a recipient bacterial cell and transforming that cell into a new bacterial species.” The media hailed the creation of “the world’s first synthetic life form.”1 Venter and colleagues at the J. Craig Venter Institute had not actually created synthetic life, although you could argue that they had reconstructed it artificially, synthesizing one chromosome of one bacterium (Mycoplasma mycoides) and transplanted it into another (Mycoides capricolum). The new bacterium was named JCVI-syn1.0, or Synthia. Whether what was created was artificial life or a new species or not, this scientific announcement reveals the remarkable extent to which scientists had become able to manipulate the genetic basis of life. The genetic technologies that allow such engineering at the level of the genome lie within the rapidly advancing field of “synthetic biology.” Synthetic biology has been described in many different ways, but in the words of the British

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Royal Society, it basically involves “the design and construction of novel artificial biological pathways, organisms and devices or the redesign of existing natural biological systems.” In simpler language, synthetic biology pioneer Rob Carlson defines the field as “bringing modern engineering principles to the design and construction of living systems.”2 The boldness of vision, and the practical ambition, of synthetic biology is arresting. In 2019, Oliver Morton wrote in The Economist of the ambition of synthetic biology: it was nothing less than “a way of controlling flows of energy on every scale from that of the smallest living cell to that of the whole living planet.” In 2009, the Emerging Risks Team of Lloyds of London wrote, “Many believe that Synthetic Biology will be one of the transformative technologies necessary to combat climate change, energy shortages, food security issues and water deficits.” Commercial applications of synthetic biology have received billions of dollars of investment globally, particularly in the United States and China. Synthetic biology is built on technologies that can determine the genetic composition of living organisms, once the gradual work of evolution. How did this come about?3 Long before there was any knowledge of the existence of DNA, people noticed that animals and plants varied, one from another. They also knew that like usually begets like—sheep with fat tails tend to produce lambs that also have fat tails (although not all offspring will resemble their parents, as Gregor Mendel showed in his peas). Some offspring exhibit features that humans found more favorable, and these offspring were encouraged to breed in hopes of getting more of those favored features. With both plants and animals, the history of domestication has been one of selective breeding—choosing parents with desired characteristics and breeding them in the hope that the trait will be passed on. Behind two hundred years of the “pedigreed” modern dog lie a thousand years of rough shaping, ten thousand years of collaboration with humans, and fifty million years of wolf-dog evolution. A dog breeder might select two animals with a particular coat color, then choose from their offspring those that best displayed the same coat color, and breed from them in turn. Eventually, such selectivity resulted in the diversity of breeds we know today— the hairless Peruvian dog, the harsh coated otter hound, or the silky Lhasa



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apso. Domestication happens with microbes as well. Penicillium molds, used in the production of bloomy rind cheeses like Camembert, have been domesticated. A variety of changes have been created, for example, shifting the volatile compounds produced from musty to cheesy. This domestication process involved changes in the expression of 356 genes.4 Human selection has perhaps been most significant in agricultural crops. For millennia, farmers have been changing the genetic makeup of plants, selecting them for size, durability, temperature and flooding tolerance, nutritional content, and countless other characteristics. The creation of modern plants like rice or wheat required hundreds of generations of selection of varieties of flowering grasses to create the familiar heavy seed heads. In the wild grasses from which wheat emerged, and in early-cultivated wheats, seeds fall off the seed head very easily (after all, the wild plant wants its seeds to be scattered). However, this trait makes harvesting difficult, so through time farmers selected for plants that did not easily scatter their seeds. In wheat, this trait is controlled by a single gene, the Q-gene. Preferential selection for particular characteristics in crops caused changes in their genomes, favoring genes that coded for the desired traits.5 Human domestication of animals and plants involved human hands laying hold of the reins of evolution and trying to steer it toward a desired outcome—a larger corncob, a head of wheat that didn’t drop its seeds, a cow that produced more milk, or a dog that could delight on a human sofa. The trouble with selective breeding is that it takes a great deal of time and is rather inefficient. In traditional breeding practices, the breeder can only work with the variation available or occasionally found in wild relatives. Flocks, fields, or gardens would be scanned to find the sweetest apple, the largest cabbage, or the chicken that laid the most eggs, to breed from them. Then the breeder has to wait until the animal or plant grows to reproductive age to harvest seeds, puppies, or lambs, pick the ones they want, then let these mature to reproductive age before starting again. With crops or animals, this could take years. Selective breeding was, even in the twentieth century, a long process often filled with failures, profoundly frustrating to crop and animal scientists and to the agricultural companies and research institutes who employed them.

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It is therefore no surprise that as the twentieth century wore on, scientists (and crop breeders in particular) began to experiment with ways to speed up the process of selection. They began to experiment with deliberately inducing mutations in DNA. Although radiation is now perhaps best known for causing genetic mutations that may in turn cause cancer, the potential of radiation to jumble genomes also seemed promising for plant and animal breeders who wanted more genetic traits to choose from. In her book Evolution Made to Order, Helen Anne Curry describes how crop breeders experimented with the X-ray machine to speed up their task. One early enthusiast gushed, “now we know that X-rays can accomplish in a few weeks what nature has been taking millions of years to do. They can speed up evolution until you get dizzy thinking about it.”6 Interest grew in the use of X-rays to create novel variety in crops and garden plants. After the Second World War, the Brookhaven National Laboratory on Long Island in the United States created an experimental field with apple trees, holly bushes, corn, oats, tomatoes, blueberries, and roses surrounding a metal pole that held aloft a piece of highly radioactive cobalt-60. A few hours of exposure a day was allowed to generate mutations in the surrounding plants. Crop breeders also started to use chemical mutagens, like colchicine, to force genetic changes (a technique instrumental in developing the seedless watermelon, among other things). After the Second World War, when the US Atomic Energy Commission became a major funder of biological research, experiments with biological applications of radiation expanded in scale and scope. Artificial mutations from X-rays and other sources helped breed minor changes in plant disease resistance, size, and flavor in over two thousand different kinds of plants, from grapefruit to chrysanthemum. Garden flower seeds were sold as “atom-blasted seeds” with all the techno-optimism of post–Second World War America. Yet neither radiation nor chemical-induced mutations lived up to their expectations.7 Radiation treatments were also applied to the control of some insect pests. Researchers conceived of the idea of using gamma-ray radiation to induce sterility in captive-raised male pest insects. If sterile males mated with females, they did not produce offspring. The idea of the so-called “sterile insect technique” was therefore to release sufficiently large numbers of irradiated males that the overall number of insects would fall. This approach



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has been applied on an area-wide basis to eradicate the New World screwworm fly (a parasite of cattle in the United States, Mexico, and Central America), some species of tsetse fly (which spreads sleeping sickness), tropical fruit flies, the pink bollworm, and the codling moth (all crop pests).8 The problem with using X-rays and chemicals to induce new and potentially useful genetic variation is that it remained unpredictable, inefficient, and costly. Most mutations produced no effect or reduced the productivity or life span of the organism. Breeders had no idea what mutations would result from the treatment, or if plants with desirable mutations would maintain the desired trait through generations. A better scientific understanding of the molecular structure of genes, and of the genome, transformed scientists’ ability to plan and bring about biological changes in domesticated species. The development of new technologies gave the power to reshape the genetic makeup of living animals and plants, doing it more rapidly, in a more targeted way, and across a much larger range of genes. Discovery of new technologies to change DNA involved many researchers over several decades. One critical step was taken by Herbert Boyer and Stanley Cohen in 1972. Boyer had discovered an enzyme (called EcoRI), which sliced up sequences of foreign DNA when a virus attacked it. Boyer and Cohen hijacked that mechanism, using EcoRI to cut and splice together DNA sequences from different organisms in a test tube. They showed that plasmids, ring-shaped lengths of DNA found in bacteria, could be physically separated from the bacterium chromosomal DNA and made to replicate independently of it. This allowed the researchers to insert genetic material from one organism into the genome of another, in such a way that the foreign DNA would replicate naturally. In effect, researchers were able to make the bacteria do something it had not done previously. Cohen and Boyer worked with a range of different organisms. In one famous experiment, they used a plasmid to introduce genes from the African clawed frog, a species commonly used in experiments, into Escherichia coli bacteria. The genes remained active in successive generations of the bacterium; they had created a reproducing bacterium that contained frog genes.9

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This new technology was not just of academic interest but was soon seen to have potential commercial use. Two aspects of this recombinant DNA technology were critically important. First, it was powerful and fast—moving a desired genetic sequence from one genome to another. Second, DNA from one species (such as a multicellular organism) could be inserted into the host of another species (typically a bacterium or a yeast). This meant that bacteria, replicating at a rate of several generations per hour, could be made to manufacture the products coded by the genes of other and slower-reproducing species, including mammals. As a result, desired biological products could be produced in the laboratory rapidly and in quantity. Commercial companies soon sprang up to exploit the technology. Two of the first products using this technology were the human hormones somatostatin and insulin. Plants and animals have much more complex immune systems than bacteria have. To work on them scientists had to learn new tricks. The first and most important species they started with was Agrobacterium tumefaciens, a naturally occurring bacterium that transfers tumor-inducing DNA-bearing plasmids into the cells of plants. It was discovered that plasmids could be altered in the lab to delete tumor-inducing genes and insert genes with desired traits. In 1974 Rudolf Jaenisch and Beatrice Mintz injected a virus found in humans and monkeys, SV40, into mouse embryos, and showed that this viral DNA was incorporated into their genomes. They had created the first genetically engineered animal.10 Frustrated with the slow pace of the work of tricking cells into taking up new DNA, scientists looked for faster ways to insert new DNA into the genome. In the 1980s, a nonbiological way of delivering new DNA into cells was developed, called a “gene gun,” or (more properly) a “biolistic particle delivery system.” It was, originally, literally a commercial air pistol, adapted to shoot tungsten pellets coated with DNA into a cell. It could be used on almost any cell, although it was most effective on large plant cells. Later, a 22-calibre nail gun was used, with a drop of DNA on the tip of a polyethylene projectile. DuPont eventually acquired rights to the gene gun in 1989 and began to develop and sell more sophisticated devices (although gene guns are still in use, often using gold pellets, which are less toxic).11 Recombinant DNA technology, the joining of DNA from two different



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species, marked a step change from traditional methods of selective breeding. For the first time it was possible to intervene directly at the genetic level to create desired change. The new methods were dramatic, relatively cheap, and fast. They found a wide range of potential applications. The animals and plants subjected to recombinant DNA technology became known as “genetically modified organisms,” or “GMOs,” to indicate that they had been altered genetically in ways that would not occur naturally by mating and/or natural recombination. The genetic variation created by GM technologies offered utterly novel prospects for scientists. Previously, artificial selection (as was the case with natural selection) had necessarily involved choices about genetic variety that already existed in a wild or domestic species. This variety (the particular order of the sequences of nucleotides in the spirals of DNA in each animal or plant) reflected longterm evolution or the previous selection decisions made in domestication and breeding. Moreover, previously, breeders could only use organisms that could breed with each other: maize with maize and camels with camels. Animals, plants, and microbes produced as a result of the new genetic technologies could be a mix of two species that could not breed. GMOs were evolutionary novelties—not directly descended from a long line of parents and offspring through evolutionary time. These novel creatures were the products of genetic creative writing—a complex neo-organism— a mix of human and natural processes and capable of breeding. If the genes introduced by the new techniques come from another variety of the same species (or a closely related species), the transfer is called “cisgenesis.” This process is essentially a more rapid and directed form of traditional breeding. It has been used to develop late-blight-resistant potatoes, apples, and cereal crops. Genetic changes made by human action in moving genes across the species boundary are called “transgenesis” (“trans” from the Latin for “across” or “beyond,” as in “across different species”). Though more dramatic and strange to humans, such movement of genes is not unprecedented in nature, as we have seen in the case of horizontal gene transfer already discussed. Transgenesis has been widely used by large agricultural biotechnology companies to develop—and in some cases market—new varieties of a number of crop species, including rice, soybean, potato, papaya, oilseeds, and many kinds of vegetables.12 Transgenesis has also been used in slightly more unnerving and even

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superficially frivolous ways. It is possible, for example, to buy engineered aquarium fish that are fluorescent, under the brand name GloFish. Genes from a jellyfish were inserted into the embryo of a zebra fish by a team in Singapore in 1999. The fish glowed, and the trait was inherited. A Texas company, Yorktown Technologies, acquired the rights and began marketing GloFish in 2003. They now sell several species in a range of colors, as do other companies outside the United States. Another fish, this time for human consumption, the genetically modified Atlantic salmon (Salmo salar) marketed in North America under the trade name AquAdvantage, is also the product of transgenesis, having genes from another fish, the ocean pout, and Pacific salmon inserted to boost rate of growth under intensive farming conditions.13 Although in some sense successful, recombinant DNA methods were undoubtedly rather crude. Gene guns shoot DNA randomly into target cells, where it can potentially enter any gene (into the nucleus, mitochondria, or any combination), and the new genetic material may be inserted multiple times in either the same or different locations in the genome. Sorting out what genetic changes have actually occurred, what new biological effects these changes cause, and how these would play out as the organism matures and breeds is quite tricky. Millions of cells would have to be tested to find the one where the insertion caused a change, and there was no reason to expect that change to be beneficial rather than harmful. GM was not as random as irradiating plants and hoping for the best, but hardly precise. After decades of research only a handful of traits reached commercial use. At the end of the twentieth century, GM technologies gave way to new approaches in genome engineering that offered far greater precision than any previous form of genetic manipulation. These techniques, conventionally referred to as “gene editing,” give scientists an unprecedented ability to design and make precise changes to DNA, independent of the cell or organism. It is this revolutionary ability that allowed the creation of Craig Venter’s “synthetic cell” described at the opening of this chapter. Gene editing is the most important technology in the field of synthetic biology. The key aspect of synthetic biology is just that, its synthesis. This element of deliberate design, with its emphasis on human intent and ac-



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tion, marks a momentous change in the relationship between humans and nature. Synthetic biology treats the genome not as something precious, created by eons of evolution, but as a raw material to be worked and shaped to meet human ends. Synthetic biology includes the modification of existing organisms, the creation of novel organisms, and work at the subcellular level, skipping down biological levels of organization to concentrate on individual chemical bases and sequences of bases. It has also become central to scientific methods for understanding biology, for example, the creation of experimental animals or plants with standardized genomes for medical research. Synthetic biology has accelerated the timeline for changing the genetic makeup of animals and plants and other organisms from millennia, centuries, or years to days or even hours.14 The year 2000 is often identified as the moment that synthetic biology decisively emerged. In that year, papers started to describe the first genetic circuits that could carry out designed functions, such as the construction of a genetic “toggle switch” that could turn gene expression on or off. Development of synthetic biology was made possible through the availability of a set of “enabling technologies” in the form of computer hardware and software. As we have seen, the approach being used is no longer that of a biologist but rather that of an engineer. Synthetic biology involves not just the application of technologies to biology, but the assumption that, in the second sentence with which Carlson starts his book, “Biology is the oldest technology.”15 Carlson, and with him synthetic biologists more generally, treat the workings of the cell, and all the complex interactions that lead to the replication of DNA, as if they were a technological system that does something. Life reproduces itself using these technologies. What the synthetic biologist does is to take a hand in directing where they want life to go and what it will do once it gets there. Thus, DNA is manipulated as standardized “parts” that consist of compatible, minimal DNA sequences that code for different biological functions. Some of these sequences might code for making proteins, and others signal for a biological process to start or stop. These “parts” can then be joined to make a “device,” a length of DNA that might start, run, or stop a biological process. These in turn can be built up into a system, a

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biological machine that can “perform” high-level tasks—for example, a bacterium coded to change color in the presence of a pollutant or engineered to break down a toxin. Synthetic biology displays its engineering heritage in its emphasis on genetic sequences imagined as a set of standardized parts. Such an approach is common to all forms of manufacturing—from the automobile to a Lego Star Wars Death Star. Standardized parts (of consistent and reliable quality) can be combined to make up a final product. If the design is changed, the same parts can be used to build something else. Genome editing is conventionally described in terms of a standardized sequence of steps. The first is to “read,” or sequence, the DNA from the original cell (determining the order of nucleotides on the DNA strand). This “reading” then allows transfer of the information from the “real” DNA to a digital description of the DNA in a computer. This digital copy is then modified to produce a new design. This modification might involve removing particular genes (or stretches of DNA), or “silencing” them by adding sets of nucleotides that do that, or cutting in genes from another organism altogether that appear to confer novel and desirable traits. The digital sequence of now-altered DNA is then sent to a DNA synthesis machine that can churn out physical DNA with nucleotides in the order specified by electronically transmitted sequence data. Finally, that DNA sequence is inserted into a living cell, which (in theory) begins operating with the new instructions, showing the desired traits. It may, of course, take many tries before this neat sequence of operations has its desired effect, but the key aim of synthetic biology is to see this as a process that can be standardized and repeated. It is precisely this abstraction of the specific choices that might be made by the gene-editing scientist and the individual changes to DNA that are intended in a standardized and repeated process—an industrial process—that is the hallmark of synthetic biology. Often referred to as “decoupling,” this approach allows synthetic biologists to program cells in a computer and have the cells made in a factory by someone else. In many ways this is decoupling the components of life from life itself. Synthetic biologists speak of a DNA sequence as a “genetic circuit,” and a cell as a “device” or “chassis.” Synthetic biology adopts a very deliberate framing of life in terms of industrial processes. Cells, genomes, and



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organisms are imagined (and treated) like machine components that can deliver useful outputs and should be tinkered with in order to improve the outcomes desired by humans. It is important to pause for a moment and reflect on the language used to describe these remarkable scientific advances, and how it makes them seem normal. DNA is often described as “the software of life.” As we have seen, the DNA code is described as being “read” and then “edited.” One enthusiastic scientist said gene editing could “be likened to editing a sentence with a word processor to delete words or correct spelling mistakes.” Once the new DNA sequence is inserted, the cell is said to be “rebooted,” like a computer awaiting a program. Synthetic biology is also described in language used only by highly trained medical staff such as surgeons. DNA is said to be cut by “molecular scissors” or a “genetic scalpel,” implying the great experience, skill, and care of those doing the work, its urgent necessity, and the implication that something is being fixed that urgently needs repair, but also that the work is done by people who know what they are doing and should be trusted.16 The implications of such metaphors are profound. The use of familiar language enables nonexperts to understand something about synthetic biology even if not any details of the science behind it. At the same time, it reveals something about the way scientists themselves visualize their interventions in the genome, as a relatively routine exercise, relatively safe. The language therefore also legitimizes and normalizes that science and obscures its messiness and the extent to which its use remains experimental.17 The key technique behind advances in synthetic biology is most commonly known as “genome editing.” There are various methods of doing this, which, as with Agrobacterium and other tools used in recombinant DNA, borrow and adapt natural systems for binding and breaking DNA. Previous editing tools included zinc fingers (engineered DNA-binding proteins) and TALENs (enzymes that can be engineered to cut specific sequences of DNA). Currently, the leading genome-editing tool arises from two discoveries about bacteria. First, bacterial DNA contains genetic sequences that are repeated. Second, the bacterium can remember viruses that had previously attacked it by storing a genetic sequence derived from that attacker’s DNA, sandwiched between repeated bacterial

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DNA sequences. This gave the bacterium adaptive immunity; using that stored genetic information, the bacterium was able to attack and cut the DNA of any virus with the same DNA sequence that came back to attack again. These genetic elements are called CRISPR, from “clustered regularly interspaced short palindromic repeats.”18 In 2012, biochemist Jennifer Doudna and colleagues showed the potential of CRISPR for genome editing. A CRISPR sequence could be used as a guide to locate what could be almost any target DNA sequence by including a copy of its particular sequence of nucleotides into a bacterium, at which point associated proteins (called Cas9, or “CRISPR-associated protein 9”) would cut the DNA at that specific point, with a high level of accuracy.19 The journal Science called CRISPR the “Breakthrough of the Year” in 2015, saying, “it’s only slightly hyperbolic to say that if scientists can dream of a genetic manipulation, CRISPR can now make it happen.” Its discovery was the result of a huge amount of work by many labora­ tories across the world. In 2020, Emmanuelle Charpentier and Jennifer Doudna received the Nobel prize for Chemistry for discovering CRISPR/ Cas9.20 The development of CRISPR is part of a progression by synthetic biologists who in the 2000s hoped that writing DNA would be a straightforward activity. This proved to be more complicated than they had hoped, and they turned to editing DNA that had already been written by evolution. Although some labs continue to pursue the de novo writing of DNA, the advent of CRISPR, and its apparent power to allow editing of  DNA, has made it the focus of most of the synthetic biology work currently being undertaken. The precision of DNA editing is increasing rapidly. In 2017 chemist David Liu developed a genetic tool that uses enzymes to make a precise rearrangement of individual atoms in the base pairs that make up the rungs in the DNA ladder. This exchanged one base pair for another, without changing anything else. Each new week brings another research development in this very rapidly evolving field. As of late 2019 six types of CRISPR cutting systems had been identified, with expectations that more will be discovered. In mid-summer 2019 Jason Chin and colleagues re-



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ported new adaptations of CRISPR that could much more accurately cut and splice millions of genes, as opposed to the hundreds of thousands previously possible. There is a great deal of work to do to replicate the findings, let alone develop them into a realistic tool, although the potential is considerable.21 CRISPR is faster and cheaper than earlier gene-editing tools, but it is far from perfect. Researchers were extremely enthusiastic as they first began to experiment with CRISPR systems. It soon became clear however that CRISPR editing suffered from a relatively high percentage of off-­ target mutations that could cause the disruption of normal genes or lead to the activation of otherwise silent genes. Research has shown, for example, that DNA breaks can lead to deletions extending over large stretches of DNA. Recent work has identified “anti-CRISPRs” that can be combined with CRISPR modules to try to limit off-target effects.22 Rapid advances in DNA editing are being accompanied by rapid advances in writing DNA at larger scales, with some discussing the potential to write entire synthetic genomes from scratch. Operating at this scale offers the opportunity to edit the entire genome. This has been done: Jonathan Venetz and colleagues of Zurich report the chemical synthesis and testing of an entire rewritten bacterial genome. Other researchers have created a synthetic bacterial genome at nearly four million bases in size.23 In engineering terms, synthetic biology is following the classic “designbuild-test-learn” cycle that has been at the heart of the development of many modern technologies, ranging from kitchen appliances to nuclear reactors, and in developing industrial manufacturing processes. In synthetic biology this cycle involves, first, proposing changes to existing genetic sequences (“design”); second, getting that changed DNA made and inserted into a cell and/or turning a gene on or off (“build”); third, seeing if the cell functions as expected from the design (“test”); and finally identifying what changes to the design are needed (“learn”). This process is conceived as a cycle, because rarely, if ever, is the first attempt successful. Scientists usually need many trials (and many failures) before something works. In a closed laboratory setting, failures can often be kept small-scale and any wider impacts contained. This is obviously harder when the object being manufactured is alive. The easy innovative

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cycles of software development, for example, provide a very poor mental model for thinking about the risks of genetic engineering out in the wildwood or the cerrado. Speeding up the design-build-test-learn cycle in theory makes it easier for researchers to learn and improve, although clearly it also has implications for possible mistakes and their impacts. The efficiency of biotechnological production processes is critical to commercial industrial production in a number of sectors, ranging from the manufacture of enzymes and biologically based materials to biopharmaceuticals or industrial feedstocks. Ginkgo Bioworks in Boston, for example, uses fifty million bases of commercially synthesized DNA per year and changes the genomes of thousands of organisms a day to help companies produce flavors, fragrances, and enzymes used in a multitude of industrial processes. Their demand is so great that they have bought a DNAsynthesis company to bring production in-house. They proudly announce on their website that they’re “making biology easier to engineer.”24 Automation is increasingly important in synthetic biology techniques. Biologists like to caution that biological systems are not entirely knowable and that it is hard to repeat experiments. But The Economist comments, “it is hard not to think that much of the unreliability is with the biologist, not the biology.” The message is clear: machines and automation are reliable where people are not. Automation and software can increase not only the speed of experimentation but also the sophistication of experimental design. Analytical machines such as mass spectrometers process the outputs of these experiments, and machine-learning software helps analyze the results.25 Increasingly, machine learning is being used to identify changes to the genomes of bacteria or yeasts or to the environment in which they are grown (nutrient supply or temperature, for example) to increase efficiency. Scaled up to the level of a factory, even marginal improvements can offer significant financial savings. Such industrial “tweaks” depend on automated technologies and high-throughput genetic synthesis. Much synthetic biology experimentation is also now automated, with racks of miniature test tubes manipulated robotically, carrying out ex­ periments to test hypotheses that are identified by computers. Specialized machines control the movement of drops of samples from one set of wells to another. The so-called “foundries,” where new organisms are assem-



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bled and replicated, have become as automated as the DNA-synthesis production systems that supply them. The easy availability of DNA constructed by automated processes pre­ sents a challenge as well as an opportunity to companies that engineer novel organisms. Humans are slow at conducting experiments compared to machines. One proposed solution is yet more automation. A genetic lab is like a factory production line: clinically clean robots handling trays of microwells, feeding in millions of tiny samples of DNA. As The Economist recently commented, “Drinking from such a firehose of DNA requires experiments designed and managed by computers.”26 Acceleration has become the norm in synthetic biology because of the remarkable and continuous fall in the cost of sequencing DNA since 2000. Costs dropped by seven orders of magnitude between 2002 and 2008 and continue to drop rapidly. The cost of sequencing DNA in the 1980s was $6,400 per base pair compared with the current cost of between $0.03 and $0.10. The price of creating new DNA has also decreased very rapidly, although not quite as fast.27 There are competing traditions to the development and deployment of standardized genetic parts. Traditionally, individual researchers secured patents on their inventions and controlled access to and use of such inventions. This contrasts with an “open source” movement in synthetic biology, mirroring that in computer software, which seeks to circumvent the powerful grip of a few giant tech companies. One of synthetic biol­ogy’s originators, and a visionary of the field, Drew Endy, supports the creation of open-access, verified libraries of genetic sequences. He suggests that the “inventors” of DNA “devices” should place in the public domain, rather than patent, their inventions. This, argues Endy, would allow the field to progress faster, without being slowed down by synthetic biologists having to negotiate rights to use patented products. Endy is one of the founders of BioBricks, which champions open access of parts and helps support the “Registry of Standard Biological Parts,” an open-access depository with over twenty thousand genetic sequences with documented functions.28 Synthetic biology is hard at work disrupting the conventional boundaries of biology itself and opening up new avenues for humans to ma­ nipulate biology to their own ends. Recent research has moved rapidly forward on “cell-free” biological synthesis. Cells, wondrous creations that

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they are, get in the way of industrial applications of synthetic biology—for example, “trapping” desired chemicals inside the cell wall, necessitating lysing (breaking down) the cell wall before being able to collect the desired product. Cells also inconveniently die, so they must be kept happy in the production setting. Production of, for example, vaccines and therapeutics, can now take place with the component parts of the cell but without the cell itself. “Cell-free” applications include biomanufacturing, and biosensing for diseases, environmental toxins and contaminants like endocrine-disrupting drugs in sewage.29 DNA is a molecule with a wide range of properties and is the subject of a considerable amount of research beyond its role in heredity, which asks how else its chemical structure could be used. For example, when Craig Venter announced his team’s creation of a synthetic cell in 2010, he also announced that they had inserted five words as “watermarks” into the DNA (including the term “Venter Institute,” in case anyone was in any doubt). The efficiency with which DNA stores information has inspired data-storage groups like the US Library of Congress to consider it for long-term storage of the books in the library—what some have termed “the ultimate hard drive.” This would involve rewriting DNA to encode and store the information in existing written texts.30 The emerging field of “DNA nanotechnology” uses the chemical structure of DNA and a system of “DNA origami” to build novel shapes. This allows DNA to carry materials such as drugs, metal nanoparticles, and proteins and deliver them to precise locations inside the human body. The hope is that such abilities will eventually allow construction of drug “nanofactories”—DNA origami that when positioned in the body can produce drugs on demand using building blocks from within a cell.31 Research is also focusing on RNA, beyond its familiar role in the making of proteins. It performs many other functions, ranging from tissue development to gene regulation. RNA is now also being developed as a geneediting tool. Unlike CRISPR, RNA editing can make temporary fixes to eliminate mutations in proteins, halt their production, or change the way they work, by altering the way genes are expressed.32 The modification of organisms using synthetic biology is also likely to play a key role in the emerging field of research on animal-machine hybrids. This is currently in its early stages and focuses on the insertion of



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electronic devices into living insects like cockroaches, beetles, or dragonflies to create living insect-microtechnology hybrids (cyborgs). Research is exploring the design and engineering of DNA as a form of biological machine making. Signs of what may be to come in this realm can be found in a tiny swimming robot designed to move like a stingray but powered by rat muscle cells genetically encoded to respond to light. It is hard to know how the conservation community will choose to regard such creatures of half-life.33 One of the weirdest but potentially most far-reaching applications of synthetic biology is “xenobiology,” literally “strange biology.” As we have explained, all forms of life on earth are based on four molecules (adenine, cytosine, thymine, and guanine), which together make up the double helix of DNA. In 2014 Floyd Romesberg at Scripps Research in California synthesized two additional bases, which had no biological analogue (prosaically, he named them “x” and “y”). These were incorporated into a cell’s DNA in such a way that they are replicated. This DNA is “artificial” on a wholly different scale from the products of conventional DNA engineering. It contains not just a sequence of bases not known from any evolved species on earth, but it contains bases that are themselves unknown.34 Such an engineered cell could be made to do things cells have never done in the history of all forms of life known on earth. For example, Floyd Romesberg and Laura Shawver have used this xenobiological approach to create a version of the cancer drug interleukin-2 that lacks its devastating side effects.35 There is no reason to stop at two synthetic bases. In 2019 Shuichi Hoshika of Firebird Biomolecular Sciences in Florida and colleagues announced that they had built DNA and RNA with eight bases (four new ones plus the original four with which DNA had evolved). This eight-­ letter DNA (called hachimoji, Japanese for “eight letters”) does everything natural DNA can do, pairing predictably and copying hachimoji RNA to direct protein synthesis. Lori Glaze of NASA Planetary Science Division observed that this new DNA structure would change thinking about the detection of extraplanetary life.36 The advent of synthetic biology is a dramatic moment in the long history of humanity’s manipulation of the natural world. Domestication made new things from old things, but it used only existing parts and was

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not as purposeful or anywhere near as fast. Synthetic biology emphasizes novelty, fabricating parts and systems that do not exist in the natural world or redesigning and fabricating those that do.37 As we have seen, existing biodiversity has evolved through the unimaginably long history of life on earth, from the first common ancestor (LUCA) to the present day. Now, however, scientists are taking apart DNA, genes, cells, and genomes and putting them back together in novel ways. How is conservation to respond to these novel forms of life, with edited or synthetic genomes? Even the most basic of life functions, photosynthesis, has caught the engineering eye of synthetic biologists. Scientists are experimenting with the insertion of enzymes from both bacteria and the human liver into plants to make photosynthesis more efficient. In synthetic biology, life is no longer even needed to make life. Petra Schwille, a biophysicist, has made the precursor of a synthetic cell using just eight ingredients: two proteins, three buffering agents, two types of fat molecules, and some chemical energy.38 Work such as this reflects an underlying sense that biology, acting on its own, is messy and inefficient, that evolution is slow and unpredictable and that there are better ways to get what humans want from nature than have traditionally been deployed. Synthetic biology therefore sets out to transform and “improve” nature with engineering. Biologist Kevin Esvelt describes synthetic biology as “sculpting evolution” and has so named his group at the Massachusetts Institute of Technology. From the conser­ vation world, Madeleine van Oppen, a coral researcher in Australia, has written persuasively of the need to “direct evolution” through biological engineering.39 The visions, ambitions, and powers offered by synthetic biology have profound implications for our thinking about nature and its conservation. These are not only conceptual, but have enormous practical significance. It matters how organisms whose genomes have been changed using genetic technologies enter the physical world, and how they interact as novel elements in its ecology. Such introductions have already begun. We turn to them, and the challenges they present, in the next chapter.

6 Synthesizing the World

Picture a small town in the heart of America. In the backyard of an ordinary house on a quiet street, a man stands over a barbecue grill. It is the weekend, and the heat of the sun is slowly easing, leaving a soupy haze, colored with the green of summer leaves and speckled with fireflies. A skateboarder rattles past on the sidewalk, and someone cracks open a beer. The familiar smell of cooking hamburgers drifts on the evening air. But let us look a little closer at our imaginary barbecue, for things are not exactly as they seem. The burgers sizzling on the grill may be made of beef, but it did not come from a cow. It is not some kind of vegetarian meat analogue, not made of textured vegetable protein or from a fungus. It does not contain cheese or peanut flour or egg whites or seaweed. It is “synthetic” or “cultured” meat, and it forms part of a new world of laboratory-based food production.1 Most attempts to create meat without killing animals have so far used what is called “cellular agriculture,” culturing meat from animal stem cells. The first cultured beefburger was grown by Mark Post, a researcher at Maastricht University, in 2013. He painstakingly assembled strips of cultured muscle tissue from stem cells taken from a donor cow and fashioned them into a burger. Costs initially looked prohibitive (the original burger cost $250,000) but have fallen so fast that cultured meat companies are

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now attracting significant investment from both philanthropists and venture capitalists. The US company Memphis Meats has been funded by entrepreneur investors such as Bill Gates and Richard Branson, as well as agribusiness giant Cargill. Mark Post’s company, Mosa Meat, raised $8.8 million on its launch in July 2018.2 In the long run, the game changer in synthetic meat is likely to be synthetic biology. Genome editing allows the production of animal proteins from completely nonanimal sources. The California company Impossible Foods already sells a range of meats made by genetically engineering yeasts to produce soy leghemoglobin heme protein at industrial scale. This adds a meaty flavor to other ingredients in their “Impossible Burger,” available in restaurants and many fast food joints.3 The reach of new genetic technologies goes far beyond synthetic meat to all parts of the human food supply chain. The barbecue chef might as easily be grilling synthetic fish: a Californian start-up, Finless Foods, grows fish fillets from lab cultures of fish cells, and genetic engineering is also being applied in the aquaculture industry. The AquAdvantage salmon, engineered by AquaBounty Technologies of Massachusetts, grows to maturity much faster than regular farmed Atlantic salmon, offering increased efficiency and shorter time to market. It went on sale in Canada in 2017, although as of 2020 it had not been approved for sale in the United States or Europe. Gene editing is being brought to the same party: CRISPR has been used successfully to edit the genomes of rainbow trout, carp, catfish, and tilapia, all major aquaculture species.4 Dairy products are another key investment target for the synthetic biologists of new biotech companies. The sour cream dip for tortilla chips, or the chocolate milkshake might well be made of cow’s milk made from genetically engineered yeast. Another Californian start-up, Perfect Day, has raised $4 million to develop “animal free milk” by fermenting yeast cells into which cow DNA has been inserted. They are rapidly moving into cheese, yogurt, and ice cream.5 Many cereals and other plant-based foods are also increasingly made from genetically engineered plants. The Flavr Savr tomato (on sale between 1994 and 1997 in the United States), its genome altered to slow ripening and extend supermarket shelf life, was the first genetically mod-



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ified (GM) crop to be licensed for human consumption. By 2015, GM crops occupied over 10 percent of the world’s arable land. The most widely grown was soybean, followed by maize and canola. Of the world’s production of soybean, 83 percent is GM, as is 29 percent of the production of maize. The United States is the world’s largest producer of GM crops, followed by Brazil, Argentina, India, and Canada. So in the United States at least, corn chips at a barbecue are likely to be made from GM corn, unless they have been purchased from an organic or other specialized producer.6 The genetic engineering revolution has also extended to clothes. What of the cotton jeans and T-shirt worn by the barbecue chef? Fully 75 percent of cotton is made from GM varieties (grown on 24 million hectares globally): Bt cotton, containing genes from the soil bacterium Bacillus thuringiensis that act as an insecticide, was first approved for commercial use in the United States in 1995. The production of other fabrics is also being influenced by advances in synthetic biology. Scientists at Utah State University have genetically engineered goats to yield the constituent proteins of spider silk and spun-silk thread. Spider silk can also be produced from genetically modified yeast in commercial quantities. The Californian company Bolt Threads has, for example, developed Microsilk, a biodegradable fiber, from engineered yeast, and is collaborating with leading fashion designers such as Stella McCartney and supplying the outdoor clothing manufacturer Patagonia.7 The same kind of revolution is being attempted in the production of leather. Another start-up, Modern Meadow, is now in trials to biofabricate leather by genetically engineering yeast to produce collagen, a protein that gives animal skin its flexibility and strength. The curing of leather is an ancient craft, traditionally using urine, dog feces, oak bark, and lime, or more recently chromium or formaldehyde. Modern Meadow makes its leather in a vat. In principle, any kind of leather can be made if the yeast DNA is programmed correctly. The Modern Meadow website urges the reader to “picture the strength of kangaroo leather but with the delicacy of snake . . . or the aesthetic of crocodile but the suppleness of lamb. Imagine if we could select the animal DNA of your choosing. Woolly mammoth leather? Not impossible.”8 Synthetic biology might even be the source of fuel for the barbecue

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grill itself. The engineering of microalgae or bacteria as a feedstock for biofuels has been a key research focus since the 1990s. Falling global oil prices have shrunk investment, but the search for a synthetic biofuel economically competitive with fossil fuels continues. In 2015, C3 BIOTECH, a start-up based at the University of Manchester, England, announced the development of a microbial biosynthetic pathway for propane and butane.9 Synthetic biology is also starting to be applied in the manufacture of high-value products like cosmetics, flavorings, scents, pharmaceuticals, and industrial chemicals from genetically engineered organisms. The California company Amyris manufactures squalane (a common ingredient in skin creams, coming—among other sources—from squalene in the livers of wild sharks) from engineered yeast cells. The Boston company Ginkgo Bioworks is engineering simple organisms to produce fragrances (for example, rose oil) and food sweeteners. It is now possible to purchase genetically modified moss in three scents (patchouli, linalool, and geranium) for use as a room freshener. In a collaboration between artists and synthetic biologists, Ginkgo Bioworks has also created perfumes from extinct trees and flowers using the DNA of herbarium specimens.10 Pharmaceuticals are also being synthesized biologically, in what is called “pharming,” altering the genome of one organism (usually a microbe or plant) to yield a biochemical product usually obtained from a more complex organism. Thus, researchers have used tobacco plants, lettuce, alfalfa, potatoes, carrots, soybeans, rice, and algae to produce therapeutic enzymes, antigens for vaccines, proteins for blood transfusions, and growth hormones, all for human medical uses.11 The classic example of such synthesis is artemisinin, traditionally extracted from the leaves of the sweet wormwood plant, grown as a crop by smallholder farmers in East Asia. Tu Youyou, a scientist at the China Academy of Chinese Medical Sciences in Beijing, shared the 2015 Nobel Prize in Physiology or Medicine for discovering that artemisinin had antimalarial properties. Scientists from the University of California Berkeley put the genes that make artemisinin into yeast, creating the possibility of industrial synthesis. Ralph Bock and colleagues at the Max Planck Institute of Molecular Plant Physiology in Germany achieved the same result by inserting the suite of genes needed to synthesize artemisinic acid into



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tobacco, yielding plants with significantly higher concentrations of the drug (although neither development found commercial success, perhaps good news for sweet wormwood farmers).12 Synthetic biology has begun to be used to turn animals into minifactories for pharmaceuticals. The targeted disruption of immunoglobulin genes and replacement with human sequences can generate transgenic chickens that express “humanized” antibodies for biopharming and functional genomics research. The genomes of other domestic animals are being hacked for the same purposes. The genomes of domestic goats have been modified to produce enzymes, anticoagulants, and anti-inflammatory proteins in their milk. Other “therapeutic proteins” are in trial, produced in milk, eggs, blood, urine, and seminal fluid of a range of domestic animals.13 The long-suffering pig is also the leading target of genome engineering to make it a factory of tissue for failing human bodies. The shortage of human organs for transplantation makes cross-species organ transplants (“xenotransplantation”) an attractive option for those who can afford it. Pig hearts have been successfully implanted into baboons, and human transplants creep ever closer. But pigs carry a set of enteroviruses that often cause transplant organs to be rejected. In 2017, a team of scientists from MIT and China used CRISPR to create a strain of pigs in which all retroviruses were inactivated, making a basis for further engineering to create safe organ and tissue resources for pig-to-human xenotransplantation.14 Behind the familiar scene of a family barbecue lies an eerie new world. Everything looks familiar, but many of its biological elements have been reengineered. Familiar products have been transformed. Yet these innovations are not science fiction. Every example (and we could have picked many more) is either already on sale or at an advanced stage of research. Some companies still use old-school GM techniques to create these new products, but these are increasingly being replaced by faster and dramatically more effective genetic technologies such as CRISPR. As a result, synthetic biology may be about to change everything that people eat, wear, and experience. Synthetic biology is not just a technical revolution, but also an economic one. Work on synthetic biology is being pushed rapidly forward

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by scientific curiosity and business investment across Europe, China, the United States, and other industrialized countries. Companies using synthetic biology tools are part of what synthetic biologist Rob Carlson calls the “bioeconomy,” the industry based on the manipulation of genes, genomes, and biological metabolism through direct modification of DNA.15 The scale and transformative potential of these new technologies is such that the World Economic Forum describes the engineering of the genomes of living organisms in terms of a “fourth industrial revolution.” It bases this claim on the way genetics, computing, robotics, artificial intelligence, 3D printing, and nanotechnology have converged. It argues that advances are blurring the distinction between the physical, digital, and biological spheres.16 Synthetic biology is a classic disruptive technology. Clayton Christensen, of the Harvard Business School, saw such disruption as the key opportunity for businesses: success was a matter of “catching the wave.” Technologies never remain fixed—they develop, mutate, stagnate, and decline, overtaken and left for dead by the rising wave of the next innovation. Success for businesses depends on endless competition, keeping up with technological change in search of efficiency of production and the reduction of costs.17 Through the nineteenth and twentieth centuries, a succession of inventions made fortunes for those who “caught the wave”: the printing press, railway, electric light, telephone, automobile, personal computer, Internet, and many more inventions have in turn shaken societies and economies to the core, changing how business is done, how people organize themselves, how landscapes are laid out, and who makes money. To eager students at leading business schools, disruption looks like a good thing, the potential magic key to corporate expansion and, potentially, market domination, and, of course, a good salary with associated benefits. But both the arrival of the new and loss of the old have huge potential significance for the global economy, for the lives and livelihoods of ordinary people worldwide, and for the environment. Investors smell the potential of the new genetic technologies to create new business opportunities, even whole new industries. Like early oilmen heading for a gusher field, they are moving in all over the world. A 2020 study by the US National Academies, Safeguarding the Bioeconomy, iden-



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tified six elements making up this new industrial push: genetically modified crops or products, bio-based industrial materials (for example, chemicals, plastics, biofuels, and agricultural feedstocks), biopharmaceuticals, biotechnology consumer products (for example, genetic testing kits and services), biotechnology business services (for example, laboratory testing and testing kits, or sequencing services), and precision medicine using biological data analysis.18 The scale of the bioeconomy is huge, although only a part of it uses synthetic biology directly. In the United States, Carlson estimated that biotech generated at least $324 billion in 2012 (over 2 percent of gross domestic product). More than forty countries now have national strategies to promote the sector. The World Economic Forum estimated that the synthetic biology industry was worth $3.9 billion in 2016 and could grow to $38.7 billion by 2020. In 2017, researchers and entrepreneurs were working on synthetic biology in over forty countries and almost 700 organizations. Between 2013 and 2017 more than 224 synthetic biology companies launched on public markets. By December 2018 over nineteen hundred patent applications had been made for CRISPR applications, about 45 percent of them in each of China and the United States.19 Although most work and investment is taking place in university or biotech labs, the synthetic biology community spans a much broader range of people, including private citizens and young people. This is possible due to synthetic biology’s use of modular pieces of DNA that are affordable and relatively available. The “maker culture” encourages “garage biology” or tinkerers and “do-it-yourself biology” (DIYbio), where tinkerers and entrepreneurs can assemble synthetic organisms without costly infrastructure. In 2020 the DIYbiosphere Project lists fifty-six community labs in the United States, Canada, Europe, India, Asia, Latin America, and Australia. Amino Labs sells kits on the Internet that will allow a curious person even without a lab to “change DNA of an organism to do fun or important things.”20 The most famous arena for young synthetic biology researchers is iGEM, the International Genetically Engineered Machine competition. Working in teams, undergraduate students use a kit of biological parts from BioBricks and compete to “build biological systems and operate them in living cells.” The 2019 iGEM Giant Jamboree in Boston, Massa-

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chusetts, attracted 353 teams and over thirty-five hundred participants representing forty-two countries. The winning high school project, from China, biosynthesized colored spider silk for use in medicine and aerospace fields.21 Synthetic biology is also being introduced to even younger age groups: on November 19, 2017, The New York Times included a special section, “CRISPR for Kids,” with a part titled “DNA Is in Every Living Thing. Get Your Scissors—It’s Time to Mess with It.” The BioBuilder Educational Foundation runs a website with projects for middle school students that include “Eau That Smell,” showing students how to make “stinky bacteria start to smell like bananas,” and another that allows students “to test a yeast that can bake vitamins into bread.”22 Although many synthetic biology innovations are still in a testing phase, or have yet to come to market, what the French researcher Mathieu Quet calls the “economy of promises” is drawing in researchers, entrepreneurs, governments, and consumers. The capacity to plan and execute designed changes in the genomes of living cells and organisms makes the synthetic biology revolution seem to turn out marvels; something that could reweave the fabric of the world’s industrial and ecological metabolism.23 Advances in synthetic biology are tweaking, remaking, imitating, and overwriting DNA in the laboratories of science and industry. Evolution, which once used to make and police the rules of biology, is being annexed, bypassed, and overwritten in the creation of novel biological products that can be brought to market. Its products have become reconceived as the raw materials of the new industrial revolution. Nature’s “biological assets” (biochemical and biomaterials) and its “biomimetic assets” (functions and processes) are, in the language of the World Economic Forum, “a significant source of economic value and future revenue.” The dazzling diversity of wild organisms on earth contains large amounts of genetic information preorganized into traits that can be identified, harnessed, and potentially modified and turned into commodities. Experimental biology has always looked to wild nature for ideas and shortcuts (CRISPR being a prime example), but in the hunt for commercial applications the search for novelty is intensifying.24 Historically, the biggest exploiters of the natural bounty of evolution



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have been plant breeders, using the genetic information in landraces (varieties grown by farmers outside the modern system), and from crop wild relatives (wild plant species closely related to domesticated crops) to develop and improve crops. The intensive scientific work of crop breeding has tended to focus on only those relatively small subsets of crop genomes linked to traits most immediately relevant to yield improvement. The longer domestication has gone on, the more uniform commercial crop genomes have become. This has become a serious problem in crops like the banana, for which the vast majority of the global crop consists of one variety, the Cavendish banana, which is vulnerable to black sigatoka virus. Most crop breeders now recognize that cultivated gene pools are shallow and need to be expanded. The hunt to expand them by capturing the genetic information in local varieties and wild plants is intensifying.25 Genetic engineering can dramatically speed up processes of plant domestication. In 2018, a Brazilian team announced de novo domestication of a wild relative of the tomato. They used CRISPR to edit six loci important for yield and productivity in domesticated tomatoes in a wild relative, creating an engineered plant with fruit three times larger and ten times more abundant than in the wild parent.26 Whatever the industrial sector, in the era of genome editing, the genetic diversity of nonhuman life has become a commodity of great potential worth. The value that the genomes of wild species offer is increasingly being recognized. Mapping the diversity of wild genomes, already seen as an urgent task by conservationists, also now attracts the biotech industry. The Earth BioGenome Project, which as we have seen proposes to sequence and catalog the genomes of all of the earth’s eukaryotic biodiversity within ten years, is justified in terms of its potential to supply genetic information that will allow the creation of synthetic fuels, new materials, new approaches to feeding the world, and new drugs. These offer important opportunities to contribute to human well-being, but they are also fields of massive commercial opportunity.27 The value of biodiversity—genes, species, and ecosystems—to biotech corporations has long been recognized. That value depends on whether genes from wild species can be patented or are regarded as a public resource. The US Supreme court ruled in 1980 that an engineered bacterium was patentable because its creation counted as “manufacture” under

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the US Patent Act of 1952. The case originated in the 1972 application by Ananda Chakrabarty, a researcher working for the US corporation General Electric, for a patent on a bacterium engineered to break down crude oil. In the United States, a patent must be novel (not previously made public), nonobvious (to someone “skilled in the art”), and useful. The creation of an engineered genome can be held to have involved “isolation and purification” into a form not available in nature, and a utility of purpose, demonstrated by linking the form of the new genome sequence to the protein it produces and the specific trait of the organism. Although a genetic sequence can be understood as a form of information (and so, like ideas or theories unable to be patented), the demonstration of a technical effect or functionality allows property rights to be claimed. The 1992 Convention on Biological Diversity’s recognized national sovereignty over genetic resources. It established principles for “bioprospecting” (searching for new species whose genes may code for useful properties), designed to help secure economic benefits for Indigenous peoples as well as for national governments in countries where the species were found. In 2010, the convention’s Nagoya Protocol set out a legal framework for the fair and equitable sharing of benefits arising out of the utilization of genetic resources, entering into force on October 12, 2014. There is currently an intense discussion about how such rules would apply if only the genetic sequence is “harvested” but not an actual specimen.28 The World Economic Forum notes that 76,274 species are listed in the global patent system, only 4 percent of all taxonomically described species and less than 1 percent of all estimated global species. To stop patenting from allowing all the benefits of evolved biodiversity to be privatized, the Forum proposes an open-access digital repository containing the genetic sequences of all Amazonian biodiversity that is run as a global public good. This Amazon Bank of Codes would record the provenance, rights, and obligations associated with all genetic material in the form of intellectual property assets on blockchain. The idea is that when value is created from these assets, smart contracts would facilitate the fair sharing of benefits with right-holders (including Indigenous peoples, whose work to identify and even domesticate species is easily forgotten). Such a model could provide economic benefits both to conserve biodiversity and to reward the people who live alongside it.29



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The connection between the natural world, with its intricate nets of evolved living diversity, and the world of the synthetic biologist’s computer, laboratory, and industrial process is close but complex. The natural world is a critical resource for biotechnology innovation. In the language of synthetic biology, evolution has generated a vast array of genetic “devices” that perform tasks that may be useful in manufacture or production. What is less widely recognized is that the connections between biotech and nature also flow back the other way. The novel genetic sequences that science and industry create can, all too easily, enter the genomes and ecosystems of the natural world. This is rarely intended and may be completely benign in its effects. It can also be profoundly problematic. The commodification of the genome is possible because of the technologies that have made it legible. In their book The Gene, Hans-Jörg Rhein­ berger and Staffan Müller-Wille argue that the Human Genome Project, and the first patent of a recombinant DNA sequence in 1980, reinforced “a conception of genes that was heavily laden with associations related to economic goods: genes appeared to be things that could be appropriated, manipulated, and alienated once again . . . exactly what one would expect from the products of a technology.”30 This transformation will affect many aspects of human lives and economies, creating winners and losers as with previous industrial innovations. But some of synthetic biology’s innovations will also have significant impacts on wild nature and biodiversity. These will mostly arise when genetically engineered species are released into the open landscape. Experiments in synthetic biology are thought of as taking place in a range of different environments. Researchers speak of “in silico” analysis (analysing genetic codes on the computer), “in vitro” experiments (experimental work done in a test tube or elsewhere outside a living organism), and “in vivo” modification, or creation of a living organism, under contained laboratory conditions. Such experiments are, and are intended to remain, confined. Genetically engineered bacteria or yeasts inside a vat, inside a laboratory or a factory, will live out their brief lives isolated from the natural world, living in artificial environments created and controlled by their owners. But not all engineered organisms will be like this. Some are being designed to live outside, in the open world. Here, experimentation takes

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place in a quite different context. Though tested in confined spaces like large enclosures, experimentation must inevitably be embedded in a world of nested and linked ecosystems, unbounded and unconfined. It takes place not just in vivo but also in eco. The most common application of synthetic biology in a free-living organism is to the human body. For the medical profession and the health industry, synthetic biology opens up an Aladdin’s cave, promising cures for genetic-based diseases, an escape from antibiotic resistance, the treatment of cancer, and many other beneficial things. Synthetic biology experiments in human health are confined, to some extent, within the human body, although human bodies leak both chemical and biological agents (as we know, for example, from the estrogen pollution in waterways caused by excretion of the metabolites of the birth control pill). Human experimentation is also closely watched and constrained by clear ethical codes and in most countries by laws (as the response to the germ-line editing of human embryos in China, the so-called “CRISPR-baby scandal” in 2019, showed very clearly).31 Yet in eco experimentation on free-living organisms is not confined to humans, and the engineering of nonhuman organisms does not share the ethical or regulatory frameworks erected around human welfare. In eco synthetic biology, by definition, takes place out in nature, in fields, woods, and oceans—the places where the problems exist that conservation is trying to save. The main sector in which this is happening is agriculture. Many applications of synthetic biology involve production under factory conditions, in closed and contained facilities. Agriculture, the commercial sector where genetic engineering has been most widely adopted, is different. Farm fields are not isolated from their surroundings. They are rather in constant communication with the ecosystems and species around, under, and above them. Moreover, crops or pasture animals with modified genomes are buffeted by the same forces of natural selection and evolution that operate in all ecosystems. Applications of genetic technologies in agriculture therefore have potential to affect wild-living organisms and their genomes, and to have direct and indirect implications for conservation. Over the last 150 years, agricultural industrialization has involved the



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use of mechanization, irrigation, artificial fertilizers, synthetic organic pesticides, crop and animal breeding, and (increasingly) automation, to increase crop yields and secure them against challenges such as drought or pest attack. As we have seen, crop breeding has been at the heart of this long agricultural revolution, the progressive adjustment of crop physiology to fit newly emerging agricultural technologies. In the modern farm regime, first developed in the United States in the 1930s, farmers bought commercial seed varieties that were bred to give consistent results, along with the fertilizer and pesticides needed to give good yields. Agricultural biotech companies became one-stop shops of industrial food production.32 In what came to be called the “Green Revolution” (for which Norman Borlaug won the Nobel Peace Prize in 1970), governments and philanthropic foundations supported programs to breed high-yielding hybrid varieties of wheat, corn (maize), and rice for the developing world. This essentially applied the models and technologies developed in US agriculture to the commercial supply of grains across the developing world. Between 1950 and 1990 the global human population rose by 110 percent but cereal production increased by 174 percent. Improved crop varieties made a significant contribution to these gains. Agricultural biotechnology companies were able to create crops that made more efficient use of financial capital, land, water, pesticides, and fertilizers. They were also able to tighten the integration of the seed and chemical sides of their business, by adapting seed genomes to the effects of their own proprietary pesticides.33 With the advent of genomics, the ability to map genomes, crop breeding became increasingly sophisticated. Gene editing, like GM before it, extended the transformation of the biological systems of agriculture to the genomic level. It allowed public and private crop-breeding organizations to speed up the search for traits such as higher yields, drought or salt tolerance, pest or disease resistance, or nutritional value.34 Novel genetic technologies in agriculture are given an important place in international policy debates about food security and climate change, for example, in the work of the Alliance for a Green Revolution in Africa, funded by the Rockefeller and Bill and Melinda Gates Foundations, and the African Orphan Crops Consortium. The latter is sequencing the ge-

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nomes of 101 neglected African crops, such as Bambara groundnut, hyacinth bean, and breadfruit. Researchers at the International Institute of Tropical Agriculture in Ibadan, Nigeria, are using genomic data to identify disease-resistant traits in African cassavas that can be combined with Asian varieties in Thailand in search of new blight-resistant varieties.35 Novel genetic technologies are also important in debates about malnutrition, for example, in efforts to improve the nutritional value of crops. This can be done by conventional breeding (for example, sweet potatoes rich in beta-carotene, a precursor of vitamin A), but transgenic methods have shown increasing promise, with early successes in crops like cereals and tomatoes.36 Golden Rice, modified to have enhanced levels of vitamin A to treat childhood blindness, is probably the best-known example of genetic engineering to improve crop nutritional value. Research began in 1982, and the findings were announced in Science in 2000. Researchers from the biotech company Syngenta introduced two new genes to Asian rice, one from maize and the other from a soil bacterium. Golden Rice proved disappointing under field conditions, and anti-GM activists destroyed field trials in the Philippines in 2013. Nonetheless, research continued, and Golden Rice is on track for licensing in the first developing country, Bangladesh.37 Genetic engineering is being used to address a number of different aspects of crop yields. A critical target is to reduce dependence on artificial nitrogen fertilizer (which is expensive and energy demanding to produce and apply, as well as causing environmental costs through runoff pollution). Two main genetic engineering approaches have been used, one that targets plant or soil microbiomes, and the other that targets the genome of the plant itself. Work on the microbiome includes the engineering of mycorrhizal species to improve nitrogen-fixing capacities. For example, Pivot Bio of California has released PROVEN, a nitrogen-producing microbial product applied to the furrow at the time of planting. Working on the plant genome, researchers at the Chinese Academy of Sciences have modified a rice gene to produce plants that use less nitrogen and have greater yields.38 A second approach to yield enhancement is directed at the core of plant biology and attempts to bring about a radical change in photosynthesis,



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the process plants use to convert sunlight into energy. Experimental work on tobacco at the US Department of Agriculture has engineered plants using genes from either bacteria or blue-green algae that increase the efficiency of photosynthesis and raise yields by 40 percent. Another effort works to reengineer rice to change the chemical pathway through which photosynthesis takes place. There are two biochemical pathways for photosynthesis in plants, called C3 and C4. The C4 pathway (used in many grasses, including maize, millet, sorghum, and sugarcane) is more efficient. Rice uses the C3 pathway. For the last two decades, researchers at the International Rice Research Institute in the Philippines and from six other institutions in five countries have been working to introduce C4 traits into rice. By doing so they hope to increase photosynthetic efficiency by 30–50 percent and also improve nitrogen use and water use efficiency. Researchers believe this may be done within the next decade.39 Another critical application of genetic technologies to crops is in combating crop pests and diseases, which are critical threats to food security and farmer income in many countries (one that is expected to become more serious with climate change). Since the 1940s, the agricultural biotechnology industry has mainly promoted the use of manufactured chemical pesticides and other compounds such as fungicides. These have been widely effective, although problematic because of their impacts on wildlife and human health.40 The long-term use of pesticides has revealed a different problem. The extreme selection imposed by their toxicity often leads many pest and disease species to evolve to be tolerant of their effects. A survey in 2017 identified 586 arthropod species, 235 fungi, and 252 weeds that had evolved resistance to at least one synthetic pesticide. The very success of the crop-­ breeding industry in selling standardized highly bred or engineered crops also means that crops over wide areas share a narrow genome, making them highly vulnerable to attack by pesticide-resistant organisms. The main industrial response to pesticide resistance has been the development of novel compounds with increased toxicity, but the resulting arms race with pests is costly and without prospect of an end. The idea of genetically engineering resistance to pests and diseases within the crop itself is therefore commercially highly attractive.41

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One of the earliest and most successful attempts to engineer pesticide resistance in a crop plant itself was in Bt maize and cotton, engineered in the 1980s to include genes from the soil bacterium Bacillus thuringiensis (Bt), which expressed proteins with insecticidal properties. Bt crops were commercially very successful, providing control of bollworm, the most important moth pest of cotton (and also of other crops, such as maize, for which Bt varieties were also developed). They went on sale in the United States in 1995, in China in 1997, in Central and South India in 2002, and across the rest of India in 2005.42 However, pests can evolve to be resistant to genetically modified traits just as easily as they can to chemical pesticides. Resistance is therefore proving a commercial as well as agronomic challenge with genetically engineered crops. From 2010, the yield benefits of Bt toxicity declined in the United States, probably because of the evolution of resistance in target species. As a result, Bt cotton still has to be sprayed with traditional insecticides for bollworm control. In India, Bt cotton initially provided good control of bollworm, but by 2009, pesticide resistance had also become a problem; pesticide use had expanded above the levels used before Bt cotton was introduced.43 To counter pest resistance to Bt toxins, Monsanto developed varieties with two toxins, and Corteva Agriscience developed a three-toxin cotton variety, WideStrike 3. Whole-genome studies of the bacterium has led to other applications of Bt toxicity aimed at the control of nematodes, mites, and ticks, and plant and animal pathogenic bacteria and fungi.44 As is now very clear, the ability to read the genomes of crops opens a library of possibilities for genetic modification to promote disease and pest resistance and control. Various synthetic biology tools, particularly CRISPR, are now being deployed to genetically engineer crops to increase resistance to disease. In bananas, for example, a gene found in red peppers has been inserted to increase resistance to banana leaf wilt. Approval has been given in the United States for farmers to deploy a genetically engineered virus to protect citrus trees from citrus greening, which has decimated orange production in Florida.45 A quite different strategy being developed against viruses and other pathogens uses RNA silencing, also known as RNA interference (RNAi). RNA silencing is a naturally occurring mechanism that can “silence,” or



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“turn off” specific targeted genes. Corn has been modified to produce an RNAi-pesticide for western corn rootworm larvae that kills them by shutting down one of their genes. In another modification of RNA, double-­ stranded RNA has been developed in Australia and deployed in a mixture of RNA and clay. When sprayed on the plant, the virus-specific RNA slowly releases from the clay, killing targeted pathogens.46 Synthetic biologists are starting to see ways to cause targeted changes in crop genomes without undertaking direct genome editing themselves. The idea is to use targeted biological agents to enter the plant’s genome and make the changes required. Researchers from Syngenta in North Carolina have shown that pollen from a corn plant modified with CRISPR can be used to carry genes into another corn plant’s cells. The pollen of these transformed plants then spreads this editing machinery to other corn varieties. The researchers also found some evidence that the CRISPR-carrying corn pollen could also edit the DNA of wheat.47 Another approach is to spray plants with double-stranded RNA (dsRNA) to trigger RNA interference in viruses and other pathogens. Using this technique, a research team from France and Finland has engineered a stable and accurate dsRNA production system in Pseudomonas syringae bacteria. This enables the broad application of dsRNA molecules to crops in greenhouses and fields to give them protection against attack by viruses and pathogens.48 In an ambitious effort that has drawn significant concern, the US Defense Advanced Research Projects Agency (DARPA) has begun funding the creation of viruses genetically engineered to suppress expression of certain defensive genes in the target organism. Viral genomes are small and relatively easy to manipulate. The DARPA work envisages viruses being dispersed by insects like pollinating bees. The simplicity of the infection process makes viral vectors an interesting alternative to the transgenic systems for the expression of foreign proteins in plants.49 At the core of environmental concern about synthetic biology is the potential for genetic and ecological impacts on nontarget organisms. Synthetic biology applications designed for industrial production are often managed within a closed factory setting. So while laboratory leakages may provide fertile material for science fiction, algae in a vat housed in a

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well-managed production facility are arguably unlikely to escape. Even if they did, they might not survive in the complicated and rich microbial communities they would encounter outside, whether in highly transformed industrial landscapes or the high-biodiversity ecosystems important to conservation.50 Yet the possibility of such an escape cannot be ruled out, and so, however unlikely, its consequences need to be understood. Where novel organisms are deliberately released into an uncontained ecosystem, such as an agricultural field, the issue becomes acute. In fields, microbes, pollen, seeds, and insects move freely into and out of crops, and with them the potential for genetically engineered genomic material to migrate into non­ target organisms, into the fields of different farmers, and into the wider environment. Even if farmers view their fields as a kind of “factory floor,” engineered for efficient food production, they cannot isolate them ecologically. Even commercial greenhouses are not normally contained systems in the laboratory sense. Containment is therefore a key problem in synthetic biology applications. As we have seen, researchers speak of a clear progression from “in silico” analysis through experimentation, “in vitro” in a cell, and “in vivo” in a living organism, to “field trials.” But a field (or an experimental plot within a field) is not an environment within which an experiment can be considered contained. A genetically engineered organism released into a field site is, to use the language of conservationists, “in the wild.” Work in the field of community genetics, as we have seen, has shown the complex ways in which genetic variation can affect ecological processes. If genetically engineered organisms are released into ecosystems, they may influence ecological interactions in ways that are not immediately obvious. It is therefore logical to expect that any genetically engineered organism released into an ecosystem will influence ecological interactions in some way. The effects could be trivial or important.51 The problem with the “escape” of a genetically engineered organism from a confined setting, be it in a laboratory, industrial production facility, or controlled field trial, is not just a question of any immediate ecological effect. The engineered organism will also be subject to natural selection, and the modification might itself be modified, making it either less



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or more fit, perhaps conferring traits that give it the potential of becoming an invasive species. Scientists and the public face a conundrum. It is not possible to know if genetically engineered organisms will have unforeseen ecological impacts without releasing them in field trials and monitoring what happens, both to the ecosystem and the flow of genes within it. And yet that very kind of experiment causes just the sort of “release” that opponents of genetic engineering fear. Even highly managed industrial agricultural fields are ecosystems, and novel genetic devices will interact with wild as well as domesticated species in unintended as well as intended ways. They will interact with microbes, fungi, plants, and animals in soil and water and adjacent habitats. They are likely to move beyond their place of application, and their genetic material will spread into other genomes and into successive generations. The effects of such a transfer on ecological interactions can be modeled but not confirmed without the field trials. The problem, as we discussed in Chapter 4, is that genes do “escape,” moving between species, a process known as genetic introgression. They also move from species with engineered genomes into nontarget organisms, a process that has been the subject of intensive scientific research. Concern about cross-species transfer of genetic material from edited to wild genomes has some basis in the high frequency of gene transfer between species in nature (referred to as “horizontal gene transfer” to distinguish it from the normal “vertical transfer” between generations). Genome sequencing is leading to the realization that genes have been widely shared across the various branches of life. As we have seen, microbes in particular are constantly exchanging genes. There is now a significant amount of research on gene flow between transgenic crops and other plants, either adjacent fields of crops or related wild species growing nearby. Gene flow within a single species (such as a crop and its wild relative) is remarkably common—flows of 10 percent are not uncommon in adjacent populations, and flows of 1 percent at a thousand meters are not unusual. Much research on genetically engineered crops relates to those developed using outmoded GM technologies. However, while the technologies have evolved, new genetic editing and old genetic modification offer the same ecological challenges in terms of gene flow from engineered organisms.52

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The tendency for genetically engineered DNA to jump beyond fields where the engineered plants have been sown can be a particular problem for organic producers, whose business models depend on being able to prove their crop is GM-free. Pollution from genetically engineered corn is a thorny problem, not least because the science of gene flow through pollen is complex and little understood. In the United States a low level of accidental contamination is permitted, but in most countries organic certification rules out most genetically engineered crops. In 2014, the Australian government revoked a farmer’s organic certification because of contamination from a neighboring farmer’s GM canola.53 Genes also move between crop plants and taxonomically related wild plants, either growing as weeds within the field, or wild in field margins or nearby ecosystems. Some important commercial crops (for example, maize, soybeans, and cotton) do not have wild relatives growing in close proximity in most of the places where they are grown. However, many other crops do, and such interspecies gene flow out of genetically engineered crop plants is widely recorded, for example, from domestic rice into wild relatives in China.54 The best researched example of gene flow between engineered cultivated crops and nearby wild relatives is the spread of engineered resistance to the broad-spectrum herbicide glyphosate. Monsanto began to market soybeans resistant to glyphosate (for which it held the patent, using the trade name Roundup) in 1996. Glyphosate kills all higher green plants (and has also been classified as a probable carcinogen to humans by the World Health Organization’s International Agency for Research on Cancer in 2015), but the genetic changes in the GM soy meant it could survive being sprayed. This made it easier for farmers to spray Roundup and kill the weeds without killing the crop. A range of so-called “Roundup Ready” GM crops, including corn, canola, alfalfa, cotton, sorghum, and wheat, were created and sold around the world. The ecological problem with “Roundup Ready” crops is that the genes conferring herbicide resistance spread out of the crop to related species among the weed community, and between wild plants within that community. This means that far from making weeds easier to kill, the genetic modification of the crop potentially makes it harder by promoting the evolution of herbicide-resistant weeds. By 2012, glyphosate-resistant weeds



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were growing on twenty-five million hectares of US cropland and had been recorded in many other countries, including Argentina, Australia, and Brazil.55 The problem of gene movement into wild plants is not confined to field crops. The US corporation ScottsMiracle-Gro engineered a herbicideresistant variety of the common plants of perennial lawns, creeping bentgrass. This is wind-pollinated and readily outcrosses with related wild species. Researchers found wild grass with the herbicide-resistant gene up to fourteen kilometers from its origin a year after the grass was planted.56 Gene flow from genetically engineered crops through pollination can carry the genes into the pollinating species. Where honeybees pollinate such crops, this has been shown to contaminate the honey supply chain. In 2012, the Mexican government gave Monsanto authorization to market GM soy, believing technical assessments that suggested that the GM plants would self-pollinate; no adverse impacts on insect pollinators were expected. However, honeybees in the Yucatán incorporated GM soy pollen from soy flowers into honey, some of which was marketed for export to countries in the European Union, where its GM content made the honey ineligible. The Mexican government withdrew Monsanto’s permit to cultivate GM soy in Mexico.57 The hybrids that result from the transfer of genes conferring pesticide resistance between engineered and wild plants normally have higher fitness compared to wild seeds and seedlings, raising the possibility of pesticideresistant “superweeds.” Research in France has shown crop-to-wild gene flow in beet crops (like sugar beet) into wild relatives as a result of hybridization events. The researchers noted that hybrid seeds could last for years in the soil seed bank. They also observed the escape of actual genetically modified seeds within the farmed landscape through spillage, and the possibility of long-distance transgene escape by human dispersal.58 However, gene introgression between domestic varieties and wild relatives does not necessarily favor the resulting hybrid. Gene introgression from planted apples is identified as a threat to the wild crab apple in Europe. Domestic apple genes reduce the fitness and genetic integrity of crab apple trees at distances up to four kilometers, potentially hindering recruitment in the wild population.59 Even if the genetic material in genetically engineered crops does not

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spread, there may still be ecological effects. Researchers talk about a “halo effect” around zones growing Bt maize and Bt cotton, in which insect pest populations are suppressed. The secondary effects of widely used engineered crops on wild species has been demonstrated by research on the famous monarch butterfly, which migrates across the United States to breed, having wintered in Mexico. In 1998, John Losey of Cornell University showed that pollen from engineered Bt corn could drift onto the milkweed plants on which monarch butterflies fed, which were growing alongside the cornfields. The US Environmental Protection Agency had not thought much about migrating monarch butterflies when they approved the Bt corn in 1995, assuming that milkweed did not grow close enough to cornfields to matter. After Losey’s paper was published in Nature in 1999, the issue hit the newspapers, and the EPA called for new research on all Bt corn products. This showed that one variety did have high levels of Bt in their pollen, while others did not. All Bt corn products except this one were relicensed in 2001.60 Single-species pest control can have unexpected effects. Where pest species interact, for example, the suppression of the population of a primary pest may increase secondary pest damage. Thus, in China, adoption of Bt cotton to suppress pink bollworm was followed by an increase in plantsucking bugs (Heteroptera). The potential for complex effects from the deployment of genetically engineered plants is especially true of impacts on soil organisms. For example, research has shown that the abundances and structures of nitrogen-transforming (ammonia-oxidizing) archaea and bacteria as well as nitrogen-fixing bacteria in the soil are significantly different when the maize host has been genetically modified.61 Genes can also move from engineered organisms into wild relatives in the ocean. Genes from genetically modified Atlantic salmon have been shown to jump into wild salmon and into brown trout. Atlantic salmon are widely farmed, both within and outside their natural range, usually in open-sided marine cages. When nets are breached, farmed salmon escape, and if they join wild salmon running up rivers to spawn, the result can be genetic introgression into wild salmon. Introgression from farmed salmon has been found in a third of Norwegian salmon rivers. It is not clear whether the standardized genomes of farmed salmon threaten the fitness of wild fish, but clearly where caged salmon are genetically engineered, these syn-



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thetic genes in farm escapes can be expected to spread into wild fish with which they have contact (unless, like the fast-growing AquAdvantage fish, all are engineered to be female and sterile). The effect on the fitness of the hybrid salmon that carry the introduced genes, and on their successors, is unknown.62 Like agronomists, forest scientists are looking to use genetic technologies to enhance growth rates and make commercial tree varieties resistant to disease or pests. Commercial forests often consist of single species stands of trees with similar genomes, making them vulnerable to pest and disease attack, just like field crops. Pesticide spraying against pests and diseases is expensive and has environmental impacts over wide areas. Trees offer a particular challenge in terms of understanding any impacts of genetic engineering, since their life spans are long and effects may take decades to emerge. Efforts to engineer tree genomes are also hampered by lack of research on the genetic basis of phenotypic traits, the complexity of forest ecosystems, and patterns of gene flow within them.63 The synthetic biology application that has drawn the greatest concern in terms of unplanned movement of DNA out of an engineered organism is the engineered gene drive. Gene drives are sequences of DNA that increase the likelihood that a given sequence of DNA will be passed from one generation to the next through sexual reproduction. Gene drives occur naturally in many species but are increasingly being created using genome editing.64 When organisms reproduce sexually the two parents each have a 50 percent chance of having their genes represented in a particular offspring. If a genetic element is attached to a gene drive, the selected gene can be passed on irrespective of its impact on fitness—thereby “driving” the traits associated with it into the genomes of all individuals in a population. Various methods have been used experimentally to create gene drives, but gene editing (especially the use of CRISPR) is proving most effective. There are some limitations on the use of gene drives. First, they only work on sexually reproducing organisms. This means, for example, that they cannot be used against agricultural pests like many aphids. Second,  they can only be used effectively in species with short generation times. They are ideally suited to species that are able to reproduce and

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disperse freely in the environment. They are well suited to insects, which reproduce quickly, although (as we will see later) they are also being developed for small mammals like mice.65 Research is being done on gene drives to control a range of agricultural pests, for example, some species of fruit fly. Focusing on plants, there is a proposal to use a gene drive to control agricultural weeds such as water hemp and Palmer amaranth.66 However, gene-drive pest control is much further advanced in the field of public health, for example, in area-wide control programs against mosquito species (such as Anopheles gambiae and Aedes aegypti). Mosquitoes are vectors of a number of serious human diseases, such as malaria, Zika, and dengue fever.67 Not all applications of genetic engineering in mosquito control involve gene drives. The biotech company Oxitec sells genetically modified male mosquitoes, branded and trademarked as “Friendly.” These carry a selflimiting gene that disrupts cell function by overproducing a protein, thereby preventing female offspring from surviving to adulthood. There have been experimental releases in the Cayman Islands, Panama, India, and Brazil. The US government is reviewing possible release in the Florida Keys.68 However, it is engineered gene drives that are currently drawing attention as a powerful and potentially highly efficient new approach to mosquito control. Two approaches are used. The first is aimed at population suppression, seeking to reduce pest densities. In this case, an inserted genetic sequence is edited to disrupt reproduction, either by knocking out the genes controlling female fertility (so they no longer lay eggs) or distorting sex chromosomes such that all progeny are male. Such gene drives and edited genes are expected to die out with the targeted insects—though natural selection might also cause them to lose effectiveness. The second approach is aimed at population replacement—modifying the genomes of all individuals in a population by replacing an existing genetic sequence with a desired one. An example would be to alter the proteins in a mosquito to which the malaria-causing Plasmodium parasite attaches, making progeny unable to transmit malaria. In this case, the engineered proteins are expected to persist in the population.69 An ambitious project called “Target Malaria,” headquartered at Im­ perial College London, proposes gene drives to reduce the population of



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malaria-transmitting mosquitoes, alongside other existing control measures. Their plan is either to skew the sex ratio of a mosquito population to produce a dominance of males, or to target genes responsible for female reproduction in order to cause sterility. Research in 2018 reported that a CRISPR gene drive that targeted the gene controlling separation of the sexes leading to female infertility spread rapidly in caged mosquitoes, reaching 100 percent of the population within seven to eleven generations. It reduced egg production to the point of total population collapse. Target Malaria’s fieldwork, focusing on altering sex ratios, is initially focused on Burkina Faso, Mali, and Uganda in sub-Saharan Africa. Controlled laboratory experiments are taking place in Africa, with plans for testing in large enclosures within laboratories in Europe.70 The possibility of releasing insects carrying engineered gene drives has been met with very significant concern from environmental groups about their potential impacts and the lack of control over the gene drive once it has been released in the wild. The whole point of a gene drive is to change many individuals of a target population without ongoing management action. Without any natural boundary to its spread, a gene drive could potentially move throughout the range of the target species. This would potentially create complex problems both of ecological impacts and of obtaining informed consent from the public.71 Scientists primarily use population and ecological modeling to assess the likely dispersal of gene-drive insects in the wild (whether around human settlements, in agricultural environments, or elsewhere). Such methods are becoming more sophisticated, but, inevitably, the behavior of genes in multispecies populations under the extreme selection pressures created by gene drives is not well understood.72 From a conservation perspective, there are several potential problems. First, the deleterious gene could spread to the target species in locations where it is an unproblematic part of the ecological community (a nature reserve adjacent to a farm field, for example). Second, crossbreeding or other processes might allow gene flow into nontarget species or populations. Third, if there are mutations in the genetic sequences attached to a drive it could spread them to both target and nontarget species. In both these cases there is the possibility that the spreading genes might turn out to be associated with undesirable traits that had been identified in trials.

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Fourth, there is the possibility that the deliberate spread of particular genes across entire populations could affect ecosystem processes.73 Not all gene-drive strategies offer the same potential to spread unsupported and to persist in nature. Malaria control researchers are most interested in “low threshold” or “nonlocalized” gene drives, in which the release of even a small number of engineered organisms (models suggest just 10 percent) leads to extensive coverage in the population. But other gene drives are being developed that are more restricted in their impacts. These include “daisy-chain” drives, designed to be self-exhausting, and “localized” or “locally fixed allele” drives, linked to unique alleles within a naturally isolated population, for example, on an island. Work is also being done on “reversal gene drives,” which overwrite the original drive and replace the genes attached to it. Such a drive might become necessary if, for example, a gene drive got out of hand or was used as part of a bioterrorist attack.74 Gene drives are still experimental, but there is already extensive debate on their deployment and potential intended and unintended impacts. There is much discussion of whether the risks offered by different kinds of gene drives are more or less acceptable, whether their use should be regulated, and what kind of limits should be placed on their deployment. It is relevant to note that gene drives are not particularly complicated to create. In 2016, students from the University of Minnesota decided to make a gene drive for their entry into the iGEM competition in Boston. They were not completely successful but came close enough to suggest the potential for independent development and release of engineered gene drives. Piers Millett, iGEM’s director of safety and security, highlighted the danger of a team working on a project that “either accidentally or deliberately resulted in something getting outside the lab.” He noted that no country had rules in place to govern how gene drives can or cannot be used in the laboratory or how they should be tested.75 Genetic modification of wild species may be hard to keep within planned limits for another reason: proliferation. The public health case is strong to eliminate disease-carrying mosquitoes. But once the technology is proven and available to pest-control companies, its use could be extended to any suitable species regarded as a pest. One recent paper on CRISPR gene drives concluded, “For the first time, we have the makings



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of a technology that could reduce or eliminate a pest population in a humane and species-specific manner.” Mosquitoes, for example, are a social nuisance even where they carry no disease threat—in the American suburban garden or a tourist destination. Control using engineered gene drives would have benefits if it reduced the amount of pesticides currently sprayed on houses, lawns, and urban waterways. But the ecological effects of removing mosquitoes from food chains are unknown. Research suggests that the ecosystem effects of removing malarial mosquitoes are limited, but larger studies are needed that include other mosquito species.76 Gene editing offers new ways to domesticate and develop crops, new approaches to pest control, and new ways to create biological products. These are the kinds of applications that are attracting venture-capital investment to biotech start-ups and fueling the advance of discovery science. But the imaginations of synthetic biologists are not constrained by such ordinariness. The biological forms and process that have not yet been produced by evolution (“the biology of what is not yet there”) offers a huge “design space” for synthetic biology to exploit. The engineering principles on which it operates: abstraction, modularity, and standardization offer the opportunity to create truly novel life-forms for novel purposes.77 Novel environmental conditions, especially those injurious to existing evolved organisms, suggest the possibility of making new forms of bio­ diversity that can survive them, or can even repair ecosystems that have become inhospitable. The artist Alexandra Daisy Ginsberg, for example, has worked with synthetic biologists to imagine novel species that might thrive in, or help manage, novel ecological futures. One creature looks like a giant slug moving on a forest floor, a “biologically-powered mobile soil bioremediation device with pH sensing and display capabilities.”78 Such mental experiments are just the beginning. Functional organisms can be imagined—and designed in silico—that diverge from any that have so far evolved. Artificial intelligence allows the definition of possible functional novel life-forms by algorithm, and in principle an automated cellbased construction tool kit could assemble them to realize living systems. The result could be a “pipeline” that “continuously outputs performant living systems” that bear little resemblance to existing organisms. These are life-forms that might be imagined to exist and could then be made to

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exist. Sam Kriegman and colleagues argue that such an approach would “pave the way to designing and deploying unique, bespoke living systems” that could serve a wide variety of functions.79 Synthetic biology could also make it possible to design living organisms that are completely novel, unlike anything that has evolved on earth. One example, as we have seen, is xenobiology, the synthesis of biologies that do not exist in nature. Only a tiny fraction of proteins that are theoretically possible occur naturally, and it is possible to identify amino acid sequences (proteins) that have a stable architecture but that do not occur in nature. This “XNA” (xeno-nucleic acid) would be “invisible” to natural biological systems. This might be useful in making a genetic “firewall” to impede exchange of genetic information with the natural world, as a biosafety tool. But it could equally wreak havoc on existing organisms if such xeno-sequences were to be transferred to other species and disrupt vital biological functions.80 Perhaps the ultimate example of a “wilder shore” for the application of synthetic biology is the idea of synthetic “terraforming.” There is interest in the synthesis of whole ecological communities—microbial communities with desirable properties. Some researchers have floated the idea that genetic engineering could be used for planetary “seeding,” with the creation of “synthetic circuit designs for earth terraformation.” The argument is that if the earth experienced catastrophic and uncontrollable environmental changes, synthetic organisms could be designed, engineered, and released to fix the problem (for example, limiting the accumulation of greenhouse gases, enhancing nitrogen fixation, or slowing down degradation in arid and semiarid ecosystems). Once a designed population of such organisms was released, the “living machines” could make copies of themselves and expand to the desired spatial and temporal scales, or even spread an engineered device across existing species. The authors of the “Earth terraformation” paper note calmly that care would be needed in any experiments, proposing to create “unescapable constraints to the engineered organisms that act as effective firewalls.”81 Synthetic biology is starting to have significant impacts on economy, society, and nature. If even a tenth of the claims made for it come to pass, its impact will expand even further. Future applications are likely to be much more ambitious and carry greater uncertainties. The release of ge-



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netically engineered organisms into the wild has many opponents. A report by Friends of the Earth questions the balance between “potential risks and hypothetical benefit.” The ETC Group (Action Group on Erosion, Technology and Concentration, an umbrella environmental organization) goes further. They argue, “The engineering of wild populations of weeds and insects to reverse resistance or make them more susceptible to chemical pesticides is a dangerous, distorted and unacceptable objective.”82 However intense the arguments are about synthetic biology and nature in general, one issue makes them particularly tricky. That is when the editing is being proposed by conservationists themselves to save a species. We explore this in the next chapter.

7 Genetic Technologies in Conservation Exploring the forested mountains of Virginia’s ancient Blue Ridge as a child, Kent paid little attention to the large, ghostly gray trunks that stood long silent among the thriving oaks. It was only later, after his father developed a minor passion for trying to replant American chestnuts, that he learned that these skeletons were all that remained of once great chestnut forests that had blanketed the eastern United States. Billions of American chestnut trees once grew from Maine to Georgia. Chestnuts were such an important part of the forests that one hill-country dweller described them as “common as the moon rise and the sun setting.”1 The American chestnut is closely related to the sweet chestnut from Europe and two related species from Asia—the Chinese chestnut and the Korean or Japanese chestnut. They are all classified in the same genus, Castanea. The chestnuts grow to over thirty meters in height and form part of the canopy layer of temperate woodlands around the Northern Hemisphere. Their trunks can grow to three meters in diameter, and their wood is dense and strong. They “coppice” freely when cut, meaning new stems grow from the stump that remains, allowing smaller timber to be cut for poles. In the fall, the trees produce abundant crops of edible nuts enclosed in prickly cases, or “burrs,” which litter the forest floor. American chestnuts played an important role in providing Native Amer-

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icans with valuable timber, bark, and food. European settlers in the eastern United States found them essential in just the same ways: they often represented the only cash crop available for many poor rural communities. The nuts, which fell from the trees in prodigious quantities, were an important food source for people, as well as for domestic and wild animals, particularly pigs, which fattened every fall on the abundant nut crop. Although heavily logged for timber through the nineteenth century, American chestnuts remained important trees in the ecological dynamics of the forests, while in rural farmlands and urban gardens and parks their size and spreading canopies provided welcome shade. All this changed in the summer of 1904. A sharp-eyed Hermann Merkel, chief forester at what is now New York’s Bronx Zoo, spotted withered leaves on one of the zoo’s chestnuts when they should have been shiny and green like on the other chestnuts in the park. Merkel’s observation was the first recorded case of a disease that would have catastrophic effects on the American chestnut across the continent.2 With dismaying rapidity chestnut trees began to die. The cause was a mystery until the curator of mycology at the New York Botanical Garden, William Alphonso Murrill, discovered the culprit was a fungus, chestnut blight canker. It had probably been introduced into the New York region in Japanese chestnut trees at the end of the 1800s. It spread with lightning speed, killing up to four billion trees from Maine to Alabama—­ everywhere chestnuts had once graced the forests. Chestnuts did not disappear without a fight on their behalf by conservationists, foresters, and the chestnut-loving public. There was a prolonged search to find disease-resistant trees across their former range, in the hope that disease-resistant saplings could be bred from this stock. Efforts were to no avail. There were even attempts to irradiate American chestnut seeds in the reactor at Brookhaven National Laboratory to try to find a way around the deadly disease (copying the methods used in crop breeding, discussed earlier). This also did not work.3 In the 1980s work began on a different approach. Scientists crossed the American chestnut with the closely related Chinese chestnut, which had evolved with the chestnut blight canker and was largely immune to its effects. Hybridized Chinese × American chestnut trees were then backcrossed with American chestnut to increase the proportion of genes of the

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American species in the new hybrid strains. The work was laborious, expensive, inefficient, and slow. Chestnuts are self-infertile, so nuts from closely related trees have low germination rates, poor rates of growth, and high mortality. Despite this, the American Chestnut Foundation, advocates, and leaders of this effort had some success. They promoted the development of “backcross breeding orchards” to breed disease-resistant hybrid trees that resembled the American chestnut. By the third generation of backcrossing of the Chinese × American hybrid with native American chestnuts, most trees showed no detectable signs of their Chinese heritage (by this stage they were 15/16 American and 1/16 Chinese, on average 94 percent American). These trees had intermediate levels of resistance to the Chestnut blight.4 The same genome-editing technologies that allowed creation of GM crops suggested a new and faster way to create disease-resistant chestnuts. In the early 2000s, William Powell and Charles Maynard of the State University of New York College of Environmental Science and Forestry used gene-editing tools to insert a gene from wheat into the forty thousand genes that make up the genome of the American chestnut. This wheat gene (also found in maize, bananas, grasses, and mosses, where it helps fight off fungal infections) produces an enzyme that breaks down a toxin called oxalic acid, which is produced by the chestnut blight fungus.5 This engineering involved transgenesis, since it inserted genetic material from a remotely related species into the American chestnut genome. The engineered American chestnut was disease-resistant and contained no Chinese chestnut genes (although it did, of course, contain the genes from the wheat). Because it added only one or two genes to the chestnut genome (out of forty thousand), the method produced trees that were genetically more similar to American chestnuts than the version crossed with the Chinese chestnut. As such, the engineered trees are closer to the species that was lost, and so, arguably, a forest of these trees would be more like the predisease forests that were lost than the interbred version.6 At present, the US government has not granted permission to plant these transgenic chestnut trees in the wild. But advocates hope that trees with the tolerance gene could be crossed with a range of surviving American chestnut trees to incorporate genetic variation naturally occurring in



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the species into disease tolerant trees, providing a broad base for ecological recovery of the species. Advocates portray the restoration of threatened “heritage” tree species as a “noble cause.”7 The American chestnut is the first genetically modified organism to be brought forward for regulatory approval in the United States for conservation purposes. Advocates of the introduction argue that the transgenic chestnut offers the best approach to restoring the American chestnut as a key species in the ecology of North American forests. It would restore lost food webs and ecological systems associated with the annual fall of nutritious nuts.8 The restoration plan for the American chestnut includes potential wild release of genetically engineered trees in close proximity to the sovereign Haudenosaunee communities of Central and Upstate New York, who had had strong historical ties to the species. Those who favor the planting of the genetically engineered trees suggest that they could contribute to cultural restoration efforts among these communities, although there is a diversity of opinions.9 However, the prospect of returning chestnuts with modified genomes to the forests of North America does not appeal to everyone. Critics express concern about the possibility of unknown and untested knock-on ecological impacts and even the possible effects on people of eating the nuts from these trees. Opponents have also voiced concern that the chestnut project is the “thin edge of the wedge” that, if approved, would allow wholesale planting of genetically modified forests for industrial harvest. In 2018, the Global Justice Ecology Project, a US-based environmental organization, dubbed the genetically engineered American chestnut tree a “Trojan horse,” saying the project was designed to help open the door to “risky commercial GE [genetically engineered] trees.” They highlighted sponsorship for the project by companies with an interest in commercial forestry, like Bayer (formerly Monsanto) and ArborGen (the largest supplier of pine seedlings for forestry projects in the United States).10 The idea of using synthetic biology for conservation purposes is both exciting and controversial. To enthusiasts, the promise of accuracy, speed, and efficiency of genome editing offers a bigger “toolbox” for conservation management. The capacity to make very precise changes to genomes

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suggests an era of “precision conservation,” and for some intractable conservation problems, synthetic biology can look like the last best hope of stopping irretrievable biodiversity loss.11 Yet the idea of making deliberate edits in the genomes of wild species makes many conservationists and nature lovers in general uneasy and, for some, downright worried. All of the problems raised by the release of genetically engineered organisms into the wild for agricultural or industrial purposes apply to conservation applications, and some are much more significant. Two kinds of conservation applications of genetic engineering have been suggested. In the first, genetic engineering is used on a species of conservation concern to improve its chances of survival. An example of this is the endangered black-footed ferret of the North American prairies. As mentioned previously, the ferret is very susceptible to the disease sylvatic plague, which periodically sweeps through prairie dog colonies. In 2018 the US Fish and Wildlife Service issued a permit to allow researchers to use genome editing to explore the possibility of giving black-footed ferrets inheritable immunity to the disease. In the second kind of application, the genomes of other species are edited to change the biological or ecological conditions for survival of the desired species, for example, the use of gene drives to control invasive species or disease.12 In either case, organisms engineered for conservation purposes are to be released in eco, free to disperse and breed in relatively undisturbed ecosystems. Indeed, for some applications (like those using engineered gene drives), this would be done with the specific intention that the altered organisms spread through that environment and modify the entire population (for example, to protect wild species against a disease or to control an invasive species). Recognizing the potential for controversy, the Convention on Biological Diversity (CBD) with its 196 member states, the United Nations Environment Programme (UNEP), and the International Union for Conservation of Nature (IUCN) with its over 1,300 members have produced analyses of the issue (Kent steered the IUCN effort). Both processes have stimulated lively debate about the science and ethics of genetic technologies in conservation.13



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One of the most prominent applications of genome editing proposed in conservation is to control invasive species. Many introduced species, alien to their new homelands, have become invasive, highly destructive of native fauna and flora. They impact adversely on biodiversity through competition, predation, or transmission of pathogens. Invasive species have affected nearly every ecosystem on earth—freshwater, marine, and terrestrial. An analysis of the IUCN Red List of Threatened Species identified alien species as a contributing cause to 25 percent of plant extinctions and 33 percent of animal extinctions. The problem is getting worse as the web of transportation that connects the world by land, air, and sea breaks down barriers to species movements.14 Nowhere is the ecological damage done by invasive species greater than on oceanic islands. Invasive species like rats and house mice are responsible for 86 percent of extinctions of island species since 1500. House mice have become established on many remote islands, and they are often disastrously destructive. Their actions can affect island ecology in a variety of ways. For example, they prey on insects and native plants, and by increasing in numbers they attract winged predators such as owls, which may in turn increase predation on native species.15 There is a sickening yet horrifically compelling video on the Internet that shows the bizarre damage that invasive species can do. A single large fluffy albatross chick sits on its nest, unmoving, as it is eaten alive by house mice. On remote Gough Island, far off the southwestern corner of Africa, critically endangered Tristan albatrosses come every year to nest, spending the rest of the year soaring over the ocean. They are long-lived (forty to sixty years), and slow to reproduce. Tristan albatrosses, like other seabirds on Gough Island, are adapted to nesting on an island so remote that it has no predators. As a consequence, albatross chicks have no defense against them. Unfortunately, at some time in the past, people (probably seal hunters) introduced house mice to Gough Island. The mice evolved in isolation and are now twice the size of normal house mice. They have become predators, preying on abundant seabird eggs and chicks. Recently they have even been shown to prey on adult albatrosses as they sit on their nests.16 The conventional approach to the control of rats, mice, and other invasive predators in confined terrain such as an island is to use traps and poison baits. Any area larger than five hectares needs poison because it is

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impractical to set enough traps and examine a high enough proportion of hiding places. The conventional approach is to mix poison with cereal or wax and broadcast it by hand or from a helicopter in such density that it to reaches and kills every rodent in the target area. This is difficult and expensive.17 As we have seen, the costs and risks of such control methods are considerable, and the poisons can kill nontarget species, so vulnerable animals have to be trapped and held in captivity while the poisoning takes place. Fear of unwanted side effects (including threats to domestic animals) can also lead local communities to oppose poisoning strategies. These problems mean that poisoning of invasive species is thought to be appropriate on only 15 percent of the islands on which endangered species are threatened by rodents. New tools that can be used on all islands are therefore a priority.18 One idea is to use an engineered gene drive to force populations of invasive species to extinction. Mice reproduce sexually and breed fast enough for an engineered gene drive to work, and oceanic islands are a relatively safe place to use a gene drive, in that the engineered mice are isolated within natural boundaries, and the variety of possible nontarget species is likely to be limited. It is also an advantage that the control method is humane, in that it does not involve any animals actually getting killed.19 House mice also make a good test bed for the development of a mammalian gene drive, because, with the laboratory rat (originally the black rat), it has the best-understood genome of all mammals by virtue of decades of selective breeding and genetic editing in medical laboratories. Research on the genetics of mice dates back to 1902, and the first mapping of part of the mouse genome was published as early as 1915. Mapping accelerated rapidly in the 1980s with advances in genomics, and increasingly large maps of the mouse genome are now available.20 Three different types of engineered gene-drive systems for mice are being investigated by an international partnership entitled Genetic Biocontrol of Invasive Rodents (GBIRd). These systems include modifying a naturally occurring gene drive (called a “t-complex”), and two engineered gene drives (CRISPR/Cas9 and CRISPR/Cpf1 [an RNA-guided form of CRISPR]). GBIRd’s objective is to inquire if it could and should create a self-limiting gene-drive modified mouse that biases future generations of



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mice to be all male (or female), thereby achieving eradication by attrition. The work is focusing on two target genes involved in development of male testes in mammals, and a chromosome that promotes solely female offspring.21 If one of these drives is introduced to captive mice and they are then released into the wild on an island, the gene drive would in theory cause these genes to spread throughout the population. If effective, the result would be that in time all mice in the population would be either male or female. Logically, this would drive the population to local extinction within a small number of generations because there could be no further breeding. To achieve this, the gene-drive mechanism would need to be strong enough to override natural selection acting against its effects, enabling it to move throughout the entire population. Mice on islands are by no means the only invasive species for which strategies based on genetic technologies are being considered. In Australia, for example, British settlers brought an ark full of alien species with them, as pets, game animals, or sport fish—to sing in their gardens or to eat agricultural pests—as well as others that came as stowaways. Invasive species are identified as the greatest threat to endemic native species in Australia. Gene-drive technologies have been proposed to control red foxes introduced from the United Kingdom and feral cats, and Tingley and colleagues have speculated about using CRISPR to target key enzymes in the pathway to the creation of the toxin in the highly invasive and poisonous cane toad, and, if successful, making the toad nontoxic.22 New Zealand is probably the country that has endured the greatest loss of native animals to invasive species. Half of the native birds of the island country have gone extinct since humans arrived, ranging from the giant moas to the laughing owl. In 2016, a consortium of government and conservationists in nongovernmental organizations (NGOs) announced the intention of making the country predator-free by 2050. The species targeted by the Predator Free 2050 program are all invasive alien mammals, including stoats, rats, and the brush-tail possum. This set of invasive species is joined by an unexpected one, wasps of two species in the genus Vespula. These social wasps can reach such densities in native forest that their biomass can be similar to, or greater than, the combined biomasses of birds, rodents, and stoats, and they compete with native animals and

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cause human injury and agricultural losses. New Zealand’s Royal Society Te Ap¯a rangi has considered the use of gene drive for all of these. For most of these species much basic biological research would be needed before a genetic intervention could be designed. But for rats, recent publication of the complete genome of the black rat opens up many possibilities for the development of gene-drive tools.23 Gene editing is also proposed to control invasive species in marine and freshwater ecosystems. As of 2008, invasive species had been reported in 84 percent of the world’s marine ecoregions. The dominant invasive groups are crustaceans, mollusks, algae, fish, and annelids (segmented worms). Researchers are using CRISPR systems to alter the genomes of a range of marine species, including diatoms, sea anemones, jellyfish, sea urchins, and the sea lamprey, some of them invasive species. Marine biologist Madeleine van Oppen, of the University of Melbourne, has suggested that an engineered gene drive might be used to reduce reproduction rates of the crown-of-thorns starfish, one of the major predators of corals in Indo-­ Pacific reefs. Their population eruptions are a major threat to corals already weakened by ocean warming and acidification.24 Many introduced plant species are also highly invasive, and synthetic biology approaches have been considered for their control. However, when compared to animals, it is less easy to deploy an engineered gene drive for plants. Many plants reproduce asexually, and gene drives only work through sexual reproduction. Gene drives also require short generation times, and many plants are long-lived; they are clearly unlikely to be useful in the control of invasive trees, for example. Despite this, attempts are being made to develop engineered gene drives to control grasses in Australia as well as agricultural weeds.25 Other applications of genome editing are also being considered for control of invasive fish. One example is the sea lamprey in the North American Great Lakes, an eel-shaped fish native to the northern and western Atlantic Ocean. It is parasitic on other fish and has a round suckerlike mouth with sharp teeth that it uses to attach itself to other fish and feed on their blood. It weakens larger fish and can kill smaller ones. Lampreys invaded the North American Great Lakes in the 1930s, changing their ecology and damaging their valuable fishery. Currently lampreys are controlled by trapping, barriers to spawning sites, and poisoning spawning and nurs-



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ery streams. Researchers are investigating an engineered gene drive against sea lamprey, genetic modification of prey species so as to poison lampreys that feed on them, and also the use of the gene silencing powers of RNA interference (RNAi). Lamprey larvae fed with RNAi under laboratory conditions suffered increasing mortality. The researchers claimed their experiment to be proof of concept for the application of this synthetic biology tool to invasive-species control.26 RNAi has also been proposed as a tool to control other invasive alien species, including insects such as ants (which are invasive in many parts of the world) and Asian long-horned beetles, which are rapidly infesting North American maples and other hardwoods. RNAi might be used to provide disease resistance, change physiology by (for example) eliminating production of a target hormone, or serve as a toxicant specific to a particular pest species.27 Conservation’s fight against wildlife disease is also starting to use techniques from the human pest-control playbook, just as invasive-species control has done. Gene drives are prominent among them. Wildlife disease is a major driver of biodiversity loss both on land and in the oceans. Diseases are natural parts of ecosystems and can influence reproduction, survival, and abundance of animal and plant populations. Disease organisms can become invasive just like rats or rabbits can, profiting by their introduction to a population that has no resistance against them. In recent decades there has been a marked increase in newly emerging diseases of wild species that have severe consequences for biodiversity. Such diseases include Ebola and three zoonotic viruses of Australasian pteropid fruit bats (the Hendra, Menangle, and Nipah viruses). Many of these viruses are “zoonotic” (meaning they also affect humans, like 2019– 2020’s disastrous coronavirus outbreak of COVID-19), which greatly increases scientific interest in addressing them. Other examples of diseases important for conservation include West Nile virus (which affects birds as well as people and is spreading in North America), and devil facial tumor disease (which affects Tasmanian devils in Australia). Not only are there now more infectious diseases of wildlife than in previous centuries, but those diseases are an increasingly significant cause of wild species decline and even extinction. Human dispersal is the commonest source of wildlife

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diseases in novel locations; it is anticipated that climate change will increase the spread of diseases.28 One example of a possible response to wildlife disease is the use of a gene drive or other genetic approaches to control mosquito-borne avian malaria, a serious threat to the surviving endemic forest birds of the remote island archipelago of Hawaii. Hawaii consists of a chain of volcanic islands, thrust up kilometers from the seafloor, separated from the nearest land by ten thousand kilometers of ocean. It has unique fauna and flora, among which the most notable are the forest birds. Of the forty-six endemic bird species on the Hawaiian archipelago at the time of European contact, only twenty-one have survived, and almost all are considered endangered. The main causes of decline are habitat loss, invasive species, and disease. A serious threat is avian malaria, an introduced disease caused by a small single-celled parasite carried by introduced mosquitoes, which enters the host’s bloodstream when a female mosquito bites. Avian malaria is getting rapidly worse on Hawaii and is a major threat to surviving endemic bird species. Conventional methods of control are not stopping bird population declines.29 The first mosquitoes probably arrived in Hawaii in the early 1800s, on board ships stopping in the islands to fill their water barrels. So far, six species have made it across the ocean, and three are particularly unwelcome. Two are a serious threat to public health and the economy: the Asian tiger mosquito and the yellow fever mosquito. The third species, the southern house mosquito, represents a different kind of threat. It is the vector for avian malaria and also avian pox virus, both diseases against which Hawaiian birds have no defense. Those forest bird species that have survived on Hawaii hang on in forest refuges high in mountains where the mosquitoes, limited by temperature and rainfall, cannot survive. Unfortunately, these refuges are shrinking, as warming temperatures mean that mosquitoes can live at higher elevations. The greatest hope of saving the remaining forest bird species seems to be to halt the spread of avian malaria and avian pox by suppressing or eradicating the non-native mosquitoes.30 A variety of versions of the sterile insect technique have been considered to save Hawaiian forest birds, including one that takes advantage of the naturally occurring bacterium Wolbachia, some species of which can



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exist only in the bodies of mosquitoes. In the lab, it is possible to introduce into mosquitoes new strains of Wolbachia that cause infertility. In theory, if such infected mosquitoes were released into the wild, the proportion of infertile mating events will reduce the size of the wild mosquito population. The US Fish and Wildlife Service, Hawaiian state agencies, and the American Bird Conservancy are exploring what it would take to implement this approach.31 The other method being discussed to control the species of mosquitoes that transmit avian malaria in Hawaii is a gene drive, just as for human malaria. A meeting of scientists and public health professionals in Hawai‘i Volcanoes National Park in 2016 reviewed this idea. Delegates recognized that the technique was relatively untested and raised some public concern. They recommended that engineered gene drives should not be deployed in Hawaii against mosquitoes until more research had been done and the public had agreed that their use was appropriate and necessary.32 Engineered gene drives seem to offer conservation many things. They provide an alternative to existing methods of controlling populations and directing the evolution of ecosystems, one that avoids many of their crude side effects. They replace the brute mechanisms of poisons, traps, or guns with a neat and apparently surgical intervention at the level of the genome. They promise to reprogram the genetic code circulating within automatically and silently. They are seductively modern and powerful. Yet they raise a number of difficult issues. First, there are questions of practicality. With invasive species such as house mice and mosquitoes, conservation stands to take advantage of decades of knowledge about their biology and genome acquired in the name of medical research. In both species, the locations on the genome that are relevant to reproduction are known, as are the genetic edits needed to create a working gene drive. So for these species, a gene drive is already well on its way to being practicable. The same is not true of other species that might be invasive or be disease threats to wildlife but that have not yet attracted research attention. Creating a tool like a gene drive for lesser-known species, for example, New Zealand’s wasps, would take significant time and money.33 Second, there are questions about the genetic and ecological impact of

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conservation gene drives, just as there are for applications in agriculture. One worry is that the engineered genes delivered by a drive may themselves have unanticipated effects. A related issue is that CRISPR editing can cause off-target mutations in the target genome, and again these may have significant and unexpected effects on the fitness of the target species. Knowledge of the links between genes and ecological traits, and of the patterns of gene flow around the target species, are important if these risks are to be understood. There is also a risk that gene flow will allow the edited genetic sequences to move out of the target organisms, potentially into another species.34 There is a range of possible unintended or indirect ecological effects of the deployment of gene drives in wild species. These extend the concerns already raised about gene drives in agricultural pest control. First, the wide distribution and proven dispersal ability of many invasive species presents a particular problem of movement of engineered genes into nontarget populations. A gene drive developed to control a species that is a pest in one place might disperse to an area where the organism is native and an integral part of the local ecology. One could imagine a mouse carrying a gene drive to control its numbers on a Pacific island hitchhiking on a pallet of bananas and getting to Europe, where Mus musculus is native. An engineered gene drive has the potential to eliminate its target species where they are wanted just as efficiently as where they are not. A second kind of impact is that removal of a species with gene-drive technology could promote unintended trophic cascades. For example, one invasive species may be replaced by another: when invasive feral domestic cats were eliminated from Ascension Island, they were replaced by black rats, which became significant major predators of breeding sooty terns. Clearly any proposed conservation use of gene drives would require careful ecological assessment. Painstaking biosecurity measures would be needed at both the genetic and ecological scales.35 As we have seen, technologies exist to minimize such risks, including the engineering of self-limiting characteristics into gene drives, or the deployment of a reversal drive to overwrite the original drive and replace its payload gene in subsequent generations. Reversal drives do not reinstate the original (“wild”) genome but add further new DNA that has the outcome of reversing the original change. Gene drives can also be targeted



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to alleles unique to an isolated population, as on a remote island. Risks of unforeseen effects from gene-drive deployment can also be reduced by extensive large-cage trials followed by field trials in isolated populations, such as those found on islands.36 Diseases caused by fungi represent a particular threat to the conservation of both animals and plants. Here, too, genetic engineering is being actively considered as a possible source of tools and methods for conservationists to use. A review in 2012 led by Matthew Fisher of Imperial College London documented the conservation challenge of fungal diseases in wildlife: they have been reported in mammals, amphibians, soft corals, bees, crayfish, and many species of plants. They are constantly showing up in new species. It appears that human activity is intensifying the impact of these diseases by dispersing infectious fungi to new locations. Fungal diseases are recognized as a particular threat to trees (as in the American chestnut blight and diseases of European ash and elm trees). An emergent and invasive fungus is also hitting hard one of the most im¯ hi‘a lehua tree. This was first obportant trees in Hawaiian forests, the ‘O served in 2014, and by 2018 it had spread to every forest district on Hawaii island.37 Fungal diseases also affect mammals. In North America, white-nose syndrome, caused by a skin fungus with the ominous name of Pseudogymnoascus destructans, has caused drastic declines in colony-nesting bats, particularly those in the genus Myotis, which overwinter colonially in caves. When infested with the white-nose fungus, hibernating bats wake up and fly about; unable to replace the energy they burn, they eventually die. The pathogen has affected bats in at least thirty-six US states and seven Canadian provinces, and some species are threatened with extinction.38 One fungal disease has drawn particular attention because of the geographical extent of its conservation impact: the global decline of amphibian species due to a fungal disease with the ungainly name of chytridiomycosis. Caused by a chytrid fungus, Batrachochytrium dendrobatidis, the disease appears to have emerged in eastern Asia around the beginning of the twentieth century. Although declines and extinctions of amphibians had been noted in a variety of places, it was not until the 1990s that the disease was recognized as a global phenomenon, and not until 1998 that

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the fungus was identified as the cause of these large losses. It has caused significant population declines and extinctions in hundreds of species, many of them endemic. Its high virulence, rapid worldwide spread, and ability to infect a wide range of taxa make the chytrid fungus an acute conservation challenge.39 To make matters worse, another species of chytrid fungus, which kills salamanders, was described in 2013. Apparently native to Asia, it was transported to Europe on animals moved for the pet trade. It kills all species of European salamanders that have been exposed to it.40 Chytrid fungi kill amphibians by attacking the keratin in their skin cells. Keratin is a structural protein that makes hair, feathers, and claws and also toughens the outer walls of cells. In amphibians, which absorb water and key electrolytes through their skin, the fungus thickens their cell walls, eventually killing them.41 There are no proven ways of curing chytridiomycosis. Laboratory experiments are being done to vaccinate frogs with antigens to the chytrid fungus or attenuated strains of it, and to augment naturally occurring antichytrid skin bacteria that seem to suppress the fungus. Both methods would be very slow and financially costly to deploy in the wild, even if they were shown to be effective. In the ongoing search for methods to control the infection, Reid Harris and Louise Rollins-Smith, leaders in the field of amphibian health, have identified two avenues by which synthetic biology might be used. Neither has yet been tested.42 The first approach is to genetically alter the pathogen using CRISPR so it loses its deadly lethality. However, it is not yet known what critical virulence factors are responsible for amphibian deaths and how strains of the disease could protect against infection. Intervention of this sort would require analysis of the fungal genome, understanding of the ecology of bacteria in the microbiome of the frog’s porous skin, the creation of a strain of bacteria with reduced virulence, and some mechanism to replace the harmful bacteria on wild frogs with the engineered bacterium. The second approach proposed by Harris and Rollins-Smith is to edit frog genomes to make them less susceptible to the disease. Three dif­ ferent parts of amphibian defense against chytrid could be considered for such interventions: innate immunity, acquired immunity, and the microbiome. Innate immunity appears to involve, among other things, secre-



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tion of antimicrobial (and apparently antifungal) peptides into the skin. It might be possible to edit the genomes of species that lack these genes so that they produce such antimicrobial peptides. The acquired immune system of an amphibian is also a potential target, but it is very complex and there is currently no apparent way to identify a set of components to target for gene editing. The final approach to defending against chytrid disease would work on the amphibian microbiome. There is strong evidence that skin bacteria in some amphibian species secrete metabolites that can protect against chytrid. The genes that produce some of these protective metabolites are known, and synthetic biology tools could be used to insert such genes into skin bacteria.43 To be useful as conservation tools, techniques such as these would have to be developed and then repeated for every species at risk, although progress would probably get much faster once a working system was identified in one species. Once the necessary changes were identified and made in frogs under experimental conditions, a viable population of engineered frogs would have to be established in the wild, although presumably the engineered frogs would spread to dominate remnant unengineered frog communities because of their greater fitness in the face of disease attack. As with all examples of synthetic biology applied to conservation, a number of questions leap to mind. First: Can this engineering be made to work? In other words, can the necessary genomic changes be identified, can the species (fungus or frog) be bred in a laboratory so that editing can be done, and will the changes have the effects on the disease that are hoped for? Second: Can the editing be made the basis of a practical conservation project, what will it cost, and where will funds come from? How many species can be treated and how long will it take? If choices have to be made between threatened species, which should be chosen? Would such a treatment result in a successfully conserved species? A third question, and it is perhaps the biggest, is what are the possible off-target effects of this genome editing? What else might be affected by the changes made to fungal or frog genome, and are there any implications of these changes being inherited by subsequent generations of frogs or fungi (for example, will the fungal genome evolve to new virulent forms in response to the evolutionary pressures imposed?); will the engineered and resurgent frogs change the ecology of their wetland homes? The

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fourth, and overarching, question is therefore will the benefits from the editing of the frogs or the fungi likely outweigh the risks, how desperate should frog conservationists feel, and will the frogs otherwise continue their slide to oblivion, or are there other possibilities? These are really tough questions that apply to many of the synthetic biology approaches we discuss. They cannot be answered without research that will take money and time, both of which are in scarce supply. It is even possible that resistance to fungal diseases will arise spontaneously. There is, for example, evidence that some frog populations are recovering following dramatic crashes in population size due to chytrid fungal infection. Recovery appears to be due to an increase in the frogs’ defenses against the disease, presumably driven by natural selection for individuals with naturally good defenses. Similarly, there is some evidence that bats are adapting to survive infection with white-nose syndrome. It is far from clear whether such adaptations will be replicated in other species or will endure.44 As conservation scientists have begun to understand the importance of the microbiomes to the survival of species like amphibians, conservation interest in the use of genetic technologies to manage them has grown. This work draws strongly on insights from human health and medicine where researchers are working on engineering so-called “smart” bacteria that have been modified to detect and eliminate diseases in the human body. They have also been engineered to act as internal sensors for signs of disease or for the production of drugs when certain diseases are detected or even to perform metabolic functions that are missing from the host due to genetic anomalies.45 One area of particular interest is the use of gene editing to manipulate the microbiomes of wild species for conservation purposes. Within the complicated bacterial communities that make up microbiomes, different species often cooperate with one another, buffering the community from environmental stress. Work on what is being called “synthetic ecology,” has begun, altering the genetics of individual microbes in order to adjust outcomes delivered by the microbial community as a whole. As we have seen, research suggests that the composition of the skin microbiome of frogs and salamanders is important to the advance of chytrid fungus, and that altering that microbiome can mitigate the impacts of the disease. Syn-



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thetic ecology approaches are also being applied to the manipulation of the microbiome in a range of other animal species.46 Plants have microbiomes, too, and work is being done, particularly with crops, to manipulate them to improve plant health and productivity. Conceivably, such approaches could be applied to species important for conservation. Sur Herrera Paredes of Stanford University and colleagues are working on designing artificial bacterial communities that can change the ways in which plants and their microbiomes respond to nutrients. It is, for example, proposed that the problem of invasive plants, such as cheatgrass, highly damaging to native grasslands in North America, might be addressed by modifying soil microbial communities to favor the reestablishment of native grasses.47 On the twenty-three-hundred-kilometer-long Great Barrier Reef off Australia, the summer of 2015–2016 was a very bad year to be a coral. Reef-building corals are amazing because they are not just animals but also plants. The coral organism, related to anemones and jellyfish, builds a hard shell—millions making up the reef—creating a place for other creatures to hide and feed. The coral organism spends the day in its coralline cave and at night emerges, extending its polyp, a fleshy protuberance, to feed. Most corals have clear bodies, and their brilliant colors come from the tiny algae that move in, photosynthesize, and produce food for the coral that hosts it. Hundreds of different algae are involved, as well as bacteria, fungi, archaea, and viruses; in fact, corals have their own microbiome.48 Synthetic biology tools are beginning to be applied in the ocean as well as on land. Record warm sea temperatures in the summer of 2015–2016 caused the Great Barrier Reef corals to expel their symbiotic algae and, as a result, it turned a ghostly white in a process called “bleaching.” Bleaching almost always kills coral, and over 60 percent of corals bleached that year. But 2015–2016 was just one of three bad years out of the past eighteen. And the Great Barrier Reef is not the only place where shallow-­ water corals are having a tough time; bleaching events have been recorded throughout the tropical oceans of the world.49 The global climate change that is driving the gradual warming of the oceans (as well as delivering occasional spikes in temperature that are particularly harmful) is also responsible for the increasing acidity of ocean

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waters. Excess carbon dioxide in the atmosphere, dissolves in the oceans, increasing the acidity. This increased acidity means that carbonate ions, which make up the shells of many organisms, are less abundant and the shells more fragile. Decreased numbers of carbonate ions in the oceans also corrode existing coral skeletons and limit the growth of new corals. Reefs become more susceptible to erosion, and if they are destroyed, so too are habitats for the myriad organisms that call coral reefs home. Some climate scenarios predict 99 percent of the world’s coral reefs will be affected by annual severe bleaching over the twenty-first century. Other projections suggest coral reefs will decline by 70–90 percent by midcentury.50 Corals can evolve through natural selection in the face of ocean warming, but projections suggest that they are highly unlikely to do so fast enough to match the speed and scale of current changes in the oceans. There is therefore increasing interest in active intervention in coral biology to increase the chances of corals surviving. A recent study commissioned by the US National Academies of Sciences, Engineering, and Medicine details a set of tools that might be used in attempts to conserve corals in the face of climate change. These include replanting bleached reefs with corals from warmer parts of the ocean, capturing eggs and sperm for future coral restoration, and cryopreservation—freezing such eggs and sperm for later use—as well as a set of tools such as shading the coral reefs, mixing cool water, and actively altering ocean acidity.51 Coral biologists and conservationists like Madeleine van Oppen are experimenting with the application of techniques of “assisted evolution,” consisting of managed selection and breeding and the use of genetic engineering to augment the capacity of corals to tolerate stress and facilitate their recovery after disturbance. Assisted evolution seeks to accelerate naturally random mutation and natural selection of corals through transplantation or assisted migration, and changes in microbial symbiont populations. This approach is essentially a version of the age-old practices of domestication, steering evolution to change the biology of a species— only this time the aim is to enable corals to survive the changing ocean. Another approach is to create hybrids between different species or between different variants within a single species that might better survive changing ocean conditions. The rapidly evolving pursuit of ways to help



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conserve corals includes working with the algal part of the coral to see if changes in the acidity of seawater can create epigenetic changes that bring increased tolerance that will persist through coral generations. Other approaches include a form of microassisted migration—this time working only with the symbiotic algae, moving them from more heat-tolerant corals to less-tolerant corals to see if they can bring greater heat tolerance abilities with them. Hannah Epstein and colleagues have proposed engineering coral microbiomes to enhance corals’ abilities to survive in the climate-altered ocean.52 Corals can and do live under physical conditions that are more extreme than those predicted for oceans changed under climate change. Marine scientist Steve Palumbi and colleagues have been working on corals off American Samoa in the Pacific. In a series of elegant experiments, they have shown that corals living in shallow water near the islands are much more tolerant of warmer temperatures than corals of the same species living in deeper water. This tolerance has a genetic and a physiological component suggesting that genetic interventions might have the potential to increase the survival of deeper-water corals in a warming ocean. Recent work at Stanford and the University of Texas has used CRISPR to edit the genome of a species of reef-building coral with the hope of identifying genes that mediate how the coral organism responds to heat and stress. Researchers also report success in altering the genome of chloroplasts inside coral-symbiotic algae.53 Synthetic biology is also being discussed as a solution to broader problems of environmental conservation and management. For example, economic logic suggests that when cheaper, better, or more attractive versions of a given product are made available, consumers will choose them. Product substitution has been identified by conservationists as a potential game changer in markets for products from wild-harvested species. One example is the market for fish oil (omega-3 oil) taken as a nutritional supplement, on which Americans spend $1.2 billion a year. Conventionally, fish oil is made by processing the millions of tons of small fish, dismissively called “forage fish.” Many of these fish come from stocks that are unregulated and harvested unsustainably. Yet such fish play important roles in ocean ecosystems, consuming phytoplankton, and supporting popula-

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tions of bigger fish, seabirds, and cetaceans. Fish do not make the omega-3 fatty acids themselves but get them from the algae they consume. In Australia, canola is now being genetically modified (by the addition of genes from an alga) to increase the level of omega-3 oil it contains. The hope is that this vegetable source of omega-3 could potentially replace the wildcaught fish-oil market, reducing fishing pressure on ocean ecosystems. It is claimed that one hectare of canola plants could provide as much omega-3 oil as ten thousand kilograms of fish. Other research is seeking to synthesize substitute fish oils from yeast, fungi, or microalgae.54 Perhaps the best example of potential direct benefits to nature through product substitution involves horseshoe crabs, those prelapsarian looking creatures that are often found tossed up on the high-tide line of beaches. In North America, Atlantic horseshoe crabs lay their eggs on the beach in huge numbers. They provide a vital food for migratory birds that feed in the tens of thousands on this energy-rich food. But the crabs also provide a substance (limulus amebocyte lysate, or LAL) that is harvested from their blood and used to detect bacteria and other contamination in human drugs and vaccines. To get the limulus amebocyte lysate, crabs are captured alive, bled, and then either returned to the ocean, turned into eel bait, or composted. All three species of horseshoe crabs in Asia, as well as the one species in North America, face overfishing, which affects the migratory birds that rely on their eggs to refuel on their long migrations north. Researchers have now engineered recombinant DNA to replicate the first reaction in the series of protein responses that signal a contaminant in the blood. They therefore have the basis of a synthetic version of LAL. It is estimated that this could reduce medical demand for horseshoe crabs by 90 percent and allow the potential recovery of the species and perhaps the migratory birds that rely on them.55 Other proposed uses of synthetic biology to derive substitutes for wild products are much less advanced in development than the replacement for horseshoe crab blood. Researchers have proposed making both synthetic rhino horn and synthetic elephant ivory. The idea is that if the synthetic horn and ivory are of identical quality and cheaper than wild sources, they could be mass-produced to undercut the market for poached products, thus removing the financial incentive for elephant and rhino



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poaching. This idea has had a decidedly mixed reception in the conservation community because it implies a continuation of the ivory and horn trade that many humanitarian as well as conservation organizations are committed to stopping. It is also thought likely that consumers of these products will reject synthesized alternatives because they see them as less efficacious than “natural” products (as they already do with products such as bear bile or tiger bone from captive animals).56 Novel genetic technologies are also spoken of in the context of ecological repair, to address the environmental harm caused by human impacts on the biosphere in the Anthropocene. Up to 4 percent of the earth’s land surface (6 billion hectares of land) is classified as degraded, meaning that the productivity of land or soil has been reduced by human activity. Little degraded land has been successfully reclaimed. Industrialization, urbanization, agricultural expansion and poor farming practices, overgrazing, and climate change have all contributed to this loss of the economic productivity of land or value for nature, or both. Impending climate change could make matters even worse over a quarter of the world’s land area.57 Genome editing is recognized as a technique that could be deployed in ecological restoration, not only (as we have seen) in the removal of in­ vasive species, but in the development of new plant varieties tailored for novel or challenging restoration contexts. In degraded drylands, communities of cyanobacteria form “biocrusts” that limit erosion, improve water retention, and restore soil carbon. In the lab cyanobacteria have been engineered to increase carbon fixation, growth, and other ecosystem functions. Such engineered microbes have the potential to help restore the many millions of hectares of degraded land.58 Human activities have not only degraded but also polluted the earth’s lands, waters, and atmosphere. Synthetic biology applications are being explored to reduce the human impact in all of these setting. Applications of synthetic biology to industrial pollution focus both on engineering processes and on cleaning up pollution once it has happened. There is, for example, interest in using synthetic biology to replace polluting chemicals in manufacturing processes, such as the use of synthetic dyes in the textile industry.59

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In terms of industrial processes, one of the most long-running discussions has been synthetic biology’s role in facilitating a move away from oil and gas as the basis of the energy economy. The replacement of fossil oil with products manufactured by microbes (conversion to what has been termed “green chemistry”) could reduce water, soil, and air pollution from oil extraction and processing, as well as reducing the climate change impacts of fossil carbon release. Research on biotech alternatives to fossil carbon is continuing, although investment has been held back by low oil prices.60 Synthetic biology is also being proposed to help in pollution cleanup. Engineered plants have been created that can degrade toxic chemicals in soil and pull toxins out of soil to prevent them leaching into ground­ water, or to signal the presence of environmental contamination through changes in growth form or color. Water treatment facilities are often not able to remove all of the pollutants present in wastewater before it is discharged into rivers or wetlands. Synthetic biology is being used to engineer microbes to help existing systems of wastewater management; for example, one effort involves the design of a bacterium that is able to degrade the antibiotic tetracycline. Synthetic biology has also been proposed as a way of addressing plastic pollution. Plastics comprise 70 percent of all the litter in the ocean and are reaching every corner of the ocean, from the deepest depths to the most remote island. Creation of biodegradable plastics offers one approach to reducing this problem.61 A number of applications of synthetic biology are being talked about in the context of responses to human-induced climate change. The release of methane (CH4) from domestic livestock (particularly cows, sheep, goats) is a critical problem. Cattle alone are said to be responsible for over 9 percent of all greenhouse gas emissions from anthropogenic sources. Methane does not last as long as CO2 in the atmosphere (less than ten years), but while it is there, its impact is much greater. Ruminants have multiple stomachs, and methane is produced by the microbes involved in digestion. Scientists have identified a set of genes that play important roles in the production of methane and might be susceptible to genetic editing. Work on pigs is further along, with researchers introducing genes for three microbial enzymes into pig genomes to help pigs metabolize feed more efficiently and produce less waste. Industrial plantations of poplar trees are also a source of a complex organic compound, isoprene, itself a



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contributor to a warming earth. Research is focusing on the use of RNA interference (RNAi) to suppress release of isoprene from plantation poplar trees.62 The most significant application of synthetic biology to climate change from a conservation perspective is the attempt to increase carbon capture and storage (to help reduce atmospheric carbon dioxide concentrations). Work is being done to augment the ability of cyanobacteria and algae to both increase their uptake and decrease their release of carbon dioxide.63 There is also extensive research to engineer the genomes of trees to increase the amount of carbon they store, by increasing growth rates. Existing natural forests provide the important ecosystem service of carbon storage, which is a strong additional argument for their conservation. However, the research focus here is on the services provided by plantations. New tree planting can lock up carbon dioxide into new timber. To maximize this ecosystem service, trees need to grow as fast as possible. Gene editing is being seen as central to the efforts of the forestry industry to maximize potential revenue from future ecosystem service payments for carbon sequestration. The biodiversity conservation impacts of such innovations are likely to be mixed. Many vulnerable species could benefit if the rate of human-­ caused climate change were slowed, but the creation of new industrial carbon plantations on existing ecosystems (either already forested or not) is likely to have negative conservation impacts. There is also a possibility that synthesized genome elements from engineered trees in carbon plantations will escape into nontarget species, or that engineered trees will spread into existing forest ecosystems and outcompete wild trees. One application of gene editing to conservation is both the most potentially novel and challenging and the weirdest. It is also, inevitably, the one that captures all the headlines: de-extinction, the re-creation of species that have already gone extinct. The return of the dead to life has long fascinated storytellers, and it features in myths like Orpheus and Eurydice, festivals like the Mexican “Dia del Muerte,” and, of course, Mary Shelley’s Frankenstein. In a modern version, the Jurassic Park films showed scientists bringing dinosaurs back to roaring life, with predictably terrifying effects.64

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The term “de-extinction” came into use in the new millennium. The idea was brought to wide public attention when the April 2013 cover of National Geographic showed a woolly mammoth and a sabre-toothed cat, with other extinct mammals, walking out of a giant test tube onto an imagined tundra landscape. An accompanying TEDx series of talks coordinated by “Revive & Restore,” a California-based nongovernmental organization, developed the theme. The series included explorations of the imminent possibility of bringing back such charismatic species as the gastric-­brooding frog (discovered in eastern Australia in the 1970s, and extinct by the 1980s due to rain forest logging), the passenger pigeon (extinct in the wild in 1900), and the woolly mammoth, which went extinct after the end of the last Ice Age.65 De-extinction is the application of synthetic biology to conservation that has excited the most comment and generated the largest number of articles, frothy excited ones, turgid ethical ones, and no small number of references to Jurassic Park, The Sorcerer’s Apprentice, and Frankenstein. In Frankenstein, life was conferred by galvanic shock. In Jurassic Park, de-­extinction required the recovery of DNA from dinosaur blood in the gut of a mosquito that had been preserved in amber. The modern version of de-extinction is based on a variety of different approaches, which range from the conventional to the novel. Gene editing is the most cutting-edge of these, and the most spectacular.66 The simplest way to bring back a lost species is to use long-established methods of selective breeding. This is called “back-breeding,” in which existing individuals of a species closely related to an extinct one are selectively bred to attain desired characteristics of the extinct species. Or, as Beth Shapiro, of the University of California Santa Cruz, puts it, to “concentrate ancestral traits that persist within a population into a single individual using selective breeding.” This requires the existence of living descendants of the desired species. Dogs, for example, have been bred to resemble wolves. Both the Sarloos wolfdog (created in 1935) and the Czechoslovakian wolfdog (created in 1955), are the result of wolf × Alsatian crosses.67 This approach was used in Germany in the 1930s to create a breed of cow that resembled the wild European auroch, which went extinct in the seventeenth century. The resulting animal (the Heck cow) has become a



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staple of rewilding projects in Europe, for example, at Oostvaardersplassen in the Netherlands. The same method is being used to re-create the quagga, a zebra subspecies, once universal on the grasslands of the South African Cape but shot to extinction in the nineteenth century. Back-breeding began in 1987, and the sequenced genome of museum specimens informed selective breeding of zebra to produce animals with partial striping (although so far without the quagga’s brown coloring).68 Another technology used in de-extinction is cloning. Cloning is a natural process in many species and involves asexual reproduction (as, for example, when the twig of a poplar tree dumped by a river flood grows into a new tree). Many cultivated plants and trees are cloned—notably fruit trees such as apples, whose cultivated varieties often cannot reproduce from seeds, cuttings of which are handed down from generation to generation of orchard and garden keepers. Animal cloning is much more recent and much more difficult. It involves the transfer of a cell nucleus from a donor individual into the unfertilized egg of a host from which the nucleus has been removed. This now-fertilized egg is then implanted in a host mother, which, hopefully, brings the offspring to term. Cellular cloning in mammals was shown to be possible when Dolly the sheep was created in 1996 at the Scottish Roslin Institute, making the cover of Time in 1997. Over the next two decades, cloning became increasingly routine as a research technique, used on chickens, mice, cattle, pigs, goats, and rabbits. It soon began to be applied to pets, first cats, and then dogs. In 2005, scientists at the Seoul National University (Korea) announced the first successful cloning of a dog: after creation and implantation of over a thousand embryos, two Afghan hound puppies were born alive. One soon died of cancer, but the other, christened Snuppy, lived for ten years before dying of the same cancer.69 Laboratories offering to clone pets now advertise on the Internet. In 2018, the actress Barbra Streisand announced that she had paid to have her ailing pet dog (a fluffy white Coton de Tulear) cloned. The two replacement dogs that resulted were genetically identical to the original, although their owner was reported to be worried that they seemed “different,” leading commentators to debate practical, emotional, and existential questions about how such clones are related to the original animal.70 Cloning extinct species requires the intact nucleus from a cell of the

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extinct species and a closely related still-alive species to act as surrogate host. As such it is of limited applicability. It is not, for example, a viable method for woolly mammoths, since even the frozen corpses now melting out of Russian permafrost lack viable eggs. But it was used to produce a living clone of the Pyrenean ibex (a subspecies of the Iberian ibex). The last of the subspecies was killed in January 2000, but tissue samples had been taken, and somatic cells were inserted into a goat as a surrogate mother. Almost all the resulting embryos died, but in July 2003, one clone was born alive, although it lived only seven minutes: its lungs were deformed.71 The use of induced pluripotent stem cells in de-extinction avoids the need for embryonic tissue or eggs—a key advantage when working with endangered species. Pluripotent stem cells—the all-purpose “jackknife” of the cellular world—are derived from skin or blood cells that have been reprogrammed genetically to become like embryonic cells. The term “pluripotent” reflects the way they can be induced to develop into many different kinds of cells. In human medicine, pluripotent stem cells have, for example, been programmed to become beta islet cells to treat diabetes, or neurons to treat neurological disorders. They are also being proposed as a technology in cellular agriculture and in de-extinction. Scientists at San Diego Zoo’s “Frozen Zoo” have used reprogramming factors from four genes to create pluripotent stem-cell lines from a range of endangered species, including antelope, birds, primates, and large cats.72 A third approach to de-extinction, and the one that has generated most of the attention, is to re-create an extinct species by reconstructing its genome using gene editing. This requires a complete genome sequence of the extinct species and of a relatively closely related living species whose genome can be edited to express the genes of the extinct species. In theory, it is possible to edit the genome of the living species by inserting into the genome the unique parts of the DNA from the extinct species to end up with an approximation of the extinct species. It may well be possible to make it resemble the original species (as the back-bred Heck cow looks very much like the extinct auroch), but the resulting organism will be a genetic hybrid—similar to the extinct species, but not genetically identical. The species that took pride of place on the cover of the National Geographic issue about de-extinction was the woolly mammoth. Mammoths are now the focus of a serious de-extinction project, led by the charis-



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matic figure of George Church of Harvard University, Beth Shapiro, and the NGO Revive & Restore. The process started with sequencing of the DNA of individual mammoths found in the permafrost of Alaska or Russia, and then reading the genome of the woolly mammoth’s closest living relative—the Asian elephant. With these two genomes in hand, it is possible to determine how mammoths differ from Asian elephants—for things like long hair and cold-adapted blood cells. Gene editing can then be used to insert these critical mammoth genes into the living cell cultures of an Asian elephant. Once that is complete, the synthesized strands of mammoth DNA can then be inserted into the egg of an Asian elephant and implanted into a host female elephant. If this fetus is brought to term and born alive, voilà, a woolly mammoth—or at least a version of one.73 The mammoth de-extinction project faces enormous technical challenges because there are about 1.4 million differences between mammoth and Asian elephant genomes. Thus far, the research team has rewritten several genes in Asian elephant cell lines, generating increasingly mammothlike cells (controlling hemoglobin, hair growth, and fat production) with each edit. Planned next steps are to reprogram fibroblasts into induced pluripotent stem cells, meaning that new cell cultures could be produced at will. Work is focusing on the regions of DNA that regulate the expression of genes and control the production of cellular proteins and pathways and editing them into the emerging woolly mammoth cell lines.74 While the science of de-extinction obviously excites those doing it, it is less clear whether de-extinction should be a priority for conservationists. Stuart Brand, cofounder of Revive & Restore, is a champion of the technology, arguing that the restoration of lost species will bring genetic diversity back into the stream of evolution. This is potentially valuable, although it will do so at a small scale, slowly, and at high cost. Paul Ehrlich, on the other hand, called de-extinction “a fascinating but dumb idea” because it could distract from more pressing issues and cost-effective conservation actions.75 Some conservationists believe that de-extinction could drive ecosystem recovery by restoring ecological function, restarting latent ecological processes, and reviving lost ecosystems. They argue that ecosystems from which species have been lost are incomplete and their functioning is impaired. In this perspective, it is not the species per se that is important,

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but what it does. In How to Clone a Mammoth, Beth Shapiro writes, “We should think of de-extinction not in terms of which life form we will bring back but what ecological interactions we would like to see restored” (emphasis in the original).76 Another argument for de-extinction is that it might increase public support for conservation. Charismatic species already exert a strong fascination for conservationists and their supporters. Charismatic extinct species might be an even greater source of wonder: as Sherkow and Greely innocently observe, “It would surely be very cool to see a woolly mammoth.”77 On the other hand, conservationists have raised a number of challenging questions about de-extinction. The most obvious one is in what sense would the individual created represent the whole lost species? A single specimen would be just as alone as the last original individual, probably (like the now-extinct thylacine—Tasmanian tiger—preserved forever on film in Hobart Zoo in 1933) in a cage. Even if the copy of the genome of one extinct individual were perfect (and this is effectively impossible), this single animal would contain just a part of the genetic diversity of the whole former species. It could not mate or breed or socialize. It might be alive but still on the cusp of re-extinction. There are also ecological questions about de-extinction. Many commentators have observed that there is no conservation point in re-creating a species if the factors that drove it to extinction have not been addressed. The scale of landscape transformation is such that many species (not least the mammoth) would struggle to find suitable habitat, especially in a warming world. It has been argued that de-extinction can only be justified in conservation terms if the re-created organisms can live outside laboratory or zoo conditions. Its aim should be the establishment of viable free-ranging populations, and the technical ability to resurrect particular species is insufficient reason to do it.78 Serious practical issues also face any project attempting de-extinction. The organisms created may well look like extinct species, but as pointed out above, they will never be genetically identical. In light of this, IUCN recommends against using the term “de-extinct,” instead suggesting “proxies of extinct species,” arguing that it avoids the moral hazard of making people think that extinction is not forever.79



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Among the practical questions about de-extinction is that of the legal status of an organism brought back through de-extinction. It is not clear whether it would be classified as the original species, either in law or in the various classificatory systems used by conservationists themselves, by different government agencies, or different governments. So, in the United States for example, would a de-extinct species be regulated as an endangered species under the US Endangered Species Act, or as a project requiring a permit under the National Environmental Policy Act, either for its possible environmental impacts, or as a GMO? Should its welfare be regulated as a wild or domesticated species? Would it be listed as endangered in the IUCN Red List, as soon as the scientist had created it? In that they will not be identical to any wild species, and their form would be the result of human ingenuity, could you patent these species? If so, any species “brought back from extinction” would be extremely valuable property. It is possible to imagine corporate investment in de-extinction, with a view to display in a private park for profit—the Jurassic Park model. It is far from clear whether such an arrangement could be accepted as a conservation project as opposed to simply a profitable public spectacle.80 Other debates around de-extinction focus on the ethics of animal welfare. Film buffs will have seen Sigourney Weaver playing the cloned Ellen Ripley in Alien Resurrection, horrified at finding a laboratory of failed human × alien hybrid clones of herself. To create Dolly the sheep, scientists transferred the nucleus from one cell into another 277 times, making twenty-nine embryos that could be implanted into surrogate ewes. Only Dolly reached term. She became famous, but her life was short. She developed arthritis in 2001 and tumors from sheep pulmonary adenomatosis in 2003. She was put to sleep, and her body put on permanent display in the National Museum of Scotland. There are clearly ethical implications to the treatment of living animals in de-extinction attempts—all of the “contenders” produced and discarded along the way. It is widely recognized that animals are “moral patients” capable of suffering, and there is a high death rate of captive wild animals in cloning.81 We opened this chapter with an example of genetic engineering in conservation deployed to engineer a disease-resistant strain of American chestnut in the hope of reestablishing the ecology of the forests that it once

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dominated. We have ended it in the fantastical world of de-extinction and the seductive idea of bringing back species that humanity has made, or helped to make, extinct. De-extinction is fascinating, complicated, expensive, and still speculative as an idea. At first sight it might look exciting from a conservation perspective, but further reflection raises many challenging questions. These range from issues of cost effectiveness and value for money, to issues of risk and ethical acceptability. The same is true, to different degrees, of all the other applications of novel genetic technologies to conservation that we have discussed in this chapter. All these applications of synthetic biology are new and untested. They awaken the same swirling mix of promise and concern as those being proposed—and implemented—in agriculture and industrial biotechnology. Behind all the scientific questions about genetic technologies lies a complex terrain of questions about whether limits should be placed on their use, how those limits should be fixed, and who should make the decisions.

8 Nature’s Future

In Margaret Atwood’s novel Oryx and Crake, the main character, Snowman, is at one point marooned on top of the walls of the wrecked compound of the bioengineering corporation RejoovenEsense, by a group of hungry pigoons. Pigoons are transgenic pigs, bioengineered to grow multiple human organs by OrganInc Farms, where Snowman’s father had once worked. They had a rapid-maturity gene spliced in, and later versions grew multiple organs that could be harvested and would regrow like a crab regrows pincers. Snowman found pigoons disquieting: “the adults were slightly frightening, with their runny noses and tiny, white-lashed pink eyes. They glanced up at him as if they saw him, really saw him, and might have plans for him later.” Pigoons had acquired enhanced intelligence along with their human genes, and in the postapocalyptic world of the novel, Snowman found himself hunted.1 Oryx and Crake updates Mary Shelley’s Frankenstein in its account of what happens when human ingenuity creates hybrids. In Frankenstein, the doctor stitches corpses together and animates them, creating a living patchwork body. In Oryx and Crake, the work of refashioning is done by geneticists. The brilliant scientist, Crake, works for one of the corporations that engineers new organisms, creating a bestiary of rakunks (mixtures of raccoons and skunks), wolvogs (with the appearance of dogs and the sav-

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ageness of wolves), and, of course, pigoons. They populate a world that is a horrific postapocalyptic synthesis of familiar and unfamiliar biologies and ecologies. The apocalypse itself resulted from a pandemic, caused by a virus created by Crake in his laboratory, and built into the BlyssPluss pill, marketed by RejoovenEsense. Like many of the fruits of genetic engineering today (particularly those in human medicine), the products sold by Margaret Atwood’s corporations were popular and found a ready market.2 The world Atwood describes was brought about in a consumer culture not entirely unlike our own. BlyssPluss was perhaps the perfect mass product for an overcrowded and polluted world. It offered protection against sexually transmitted diseases, and promised heightened energy, unlimited libido, and a sense of well-being and prolonged youth. But it also made its users sterile, combining mass gratification and population control. Unfortunately, it eventually killed everyone who took it, leaving only Snowman (Crake’s former friend), and the Crakers, near-humans whose DNA had been genetically engineered by splicing in characteristics of other animals to make them docile. At the end of Mary Shelley’s novel, Frankenstein’s creation was hunted down by its creator, bent on vengeance. The so-called “daemon” was last seen on a block of polar ice, being swept away into the darkness. The voice of the narrator in the closing chapter frames the story as a horrific but finished episode in a world that remained normal. By contrast, Atwood’s pigoons and other literary creations remain thriving and dangerous at the end of the book. Her imagined world is one that has been permanently changed. The corporate experiments are alive and breeding, their genomes adapting and evolving in a permanent genetic dystopia. Oryx and Crake and Frankenstein are part of a rich seam of stories about the power of biotechnology to disrupt familiar categories in nature. One might compare them with the uncanny sinister life-forms of Area X, in Jeff VanderMeer’s Annihilation. Strangeness and hybridity are at the heart of this brand of fiction. The stories work because the same strangeness pervades reactions to technological change in wider society, particularly where it seems to blur or cross recognized boundaries. As biotechnology, and the ability to edit genes and transform the bodies of living organisms, has advanced, it has come to epitomize unease at such transgressions.3 But there is a second element to these science fictions, and this is the



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question of authority and power. Who does the experiments that can remake life? Frankenstein was independently wealthy, a man who had no need of a research institution with its ethics panels, or a funding body, or even colleagues. In Margaret Atwood’s world, it was corporations (the wonderfully named RejoovenEsense, OrganInc Farms and HelthWyzer) that ruled. In the absence of meaningful government, their wealth and power were unconstrained, their work unregulated. They were free to design, build, and market whatever they wished, to a hungry, desperate, and dysfunctional world. Only the fear of corporate espionage limited their ambition and reach. Atwood is by no means alone in her dystopian view of unbridled markets and corporate power in the context of genetic engineering. Paolo Bacigalupi’s The Windup Girl describes a future Bangkok, hunkered behind high seawalls in a world crippled by climate change and ecological collapse. Global food production is controlled by a series of giant corporate “calorie companies” whose genetically engineered crops are threatened by blights and diseases created by the very “genehacking” used to create them. Thailand holds the only remaining seedbank of plants that have not been engineered, and bans import of all foreign genetic material. The story turns on the AgriGen corporation’s attempt to subvert their security and get access to the seeds and the genes within them that they need to fix past mistakes.4 Fiction writers also offer a pessimistic view of one possible future of engineered wild species. In his novel Tears of the Trufflepig, Fernando Flores follows the consequences of a new technology, vaguely referred to as “filtering,” created during a time of food shortage to grow food. Syndicates along the United States–Mexico border find ways to use filtering to bring back extinct and rare animals so that they can cook and serve them in pricey dinner parties or use their feathers and fur as luxury items: de-extinction, but hardly for conservation purposes.5 The scientific advances that allow the editing of genomes and the creation of novel forms of life, described in science fiction works like those, are, as we have seen, starting to have widespread impact in fields such as biotechnology and agriculture. Scientists can now edit genes and are doing so in the laboratories of government research agencies, universities, and private corporations, in classrooms and in garages around the world.

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Meanwhile, genetic engineering offers conservationists the dizzy prospect of the power to counteract relentless biodiversity loss by saving species, replenishing depleted genomes, making coral reefs resistant to a warming ocean, restoring degraded environments, and even, possibly, bringing extinct species back to life. Who could be against these things? But the choice to adopt genetic technologies is less simple than it might seem. Gene editing also carries risks for nature. It is already being used on wild species (mostly so-called pest species), in breeding crops, and in novel industrial processes. Engineered organisms are already loose in agricultural fields. Who fully understands what impacts they, or those engineered for conservation, might have, and how should such uses be approved? Whatever decisions come to be made about the use of genetic technologies in conservation, they will be shaped by public attitudes forged in wider debates stimulated by developments in agriculture, biotechnology, and human health. For example, the rise of the COVID-19 pandemic has led some, like The Economist, to predict that as a result the world will more quickly adopt new technologies, including gene editing. Such debates, in their turn, are often proxies for wider issues, for example, about food security, social justice, food safety, the environmental impact of farming systems, the balance between intellectual property protection and benefit sharing, or the power of multinational companies.6 Behind concerns about the risks of gene editing to wild nature lies unease about the human power to rewrite the codes that underpin the familiar features of life, and fear that these codes cannot be changed without unpredictable harm. Yet the scale at which these technologies are already being used means that they cannot, like Aladdin’s genie, simply be put back in the lamp. It is much too late for that. In 2017, the US National Academies classified categories of biotechnology products likely to be developed in the United States in terms of the extent to which they have had precedents within the US regulatory system. The first consists of things that are broadly similar to existing biotech products, such as genetically engineered crops and fermentationbased production of small molecules, enzymes, or other biochemicals. Many of these are already approved and commercially available. The second category included innovations under development that expand on



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familiar species and genetic pathways, for which there are well-established approaches to assessing risk—including products derived from animal cell culture (such as no-cow meat or leather) and plants for bioremediation, decoration, or other environmental or consumer use. These products, too, find precedents in US regulation. The third and fourth categories are different. They are “first-of-their-kind” products and have no precursors. The third category consists of products that use multiple organisms in complex microbial communities, such as microbiome engineering and synthetic consortia for bioremediation or bio-mining applications. The fourth and last category involves the use of rapid design-build-test-learn cycles to design novel genetic pathways in diverse species, including open release into the environment of organisms intended to modify populations of natural species. Examples here include some of those discussed in this book, such as genetically engineered mosquitoes to fight malaria and genetically engineered microorganisms. All four categories lie within the realm of both practical science and commercial investment.7 It goes without saying that not all parties are convinced by promises made about the benefits of biotechnology innovations in any of these categories. Indeed, applications such as engineered gene drives (with their ability to spread on their own) alarm as many people as they excite. The technology media and opposition groups buzz around new announcements like wasps around a jam pot. The questions of whether genetic engineering poses ecological risks, and the extent to which these can be traded off against benefits, is a critical one. It is debated in a world where data are scarce but many people have already taken sides. Over the last decade, as we have developed our thinking and this book, we have felt the heat of debates about synthetic biology on several occasions. At various times one or other of us have been accused of building a Trojan horse to disguise the interests of commercial synthetic biology for even discussing applications within conservation. The meeting that we organized in Cambridge with the late Professor Georgina Mace back in 2013, “How Will Synthetic Biology and Conservation Shape the Future of Nature?,” brought people from the conservation and synthetic biology communities together for the first time to talk about how new genetic technologies might affect nature, for good or ill. The day before the meeting started, we received an e-mail from someone who had heard about it but

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was not attending. They put it to us that the genetic technologies we were to discuss would have unintended consequences for biodiversity that were just as destructive as the problems they were supposed to deal with. We were, they said, like cocaine addicts, stubbornly believing that the impacts of drug abuse needed to be countered with more drugs. They saw no adequate mechanism to manage the powers synthetic biology could unleash. This note captures the core of many concerns that we have heard (and have continued to worry about ourselves) concerning conservation uses of synthetic biology.8 Between 2017 and 2019, Kent led an evidence-based assessment of potential synthetic biology applications to conservation for IUCN, the International Union for Conservation of Nature. IUCN’s thirteen hundred members (governmental, nongovernmental, and Indigenous organizations drawn from over 170 countries) commissioned the assessment at the 2016 World Conservation Congress in Hawaii, as a precursor to the development of a policy on conservation applications. The Technical Subgroup on Synthetic Biology and Biodiversity Conservation consisted of seventeen people, drawn from universities, research institutes, governments, NGOs, and IUCN itself, based in 10 countries, plus a number of other contributing authors. It was charged with examining the impacts of the production and use of products resulting from synthetic biology on the conservation and sustainable use of biodiversity. The assessment ran to 166 pages and was put up on the IUCN website in draft form for comment and review in 2018. Over five hundred comments were received and responded to, with the responses posted online. The final version was published in 2019. It offered a summary of all available evidence relating to the possible application of synthetic biology to conservation and reviewed other applications that might have conservation impacts. It included a set of case studies and also addressed issues of the governance of the new technologies and the way “evidence” (like that used in the assessment) could be understood and used.9 The report came in for a substantial amount of criticism, in the first instance mainly from nongovernmental organizations that were not members of IUCN. Critics dismissed the report as biased because the IUCN Task Force had included people who were working synthetic biologists. Kent presented the draft report to a workshop at the biennial meeting of



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the Convention on Biological Diversity in Egypt. Afterward, a lobbyist said to him that the composition of the report writing team was its Achilles’ heel, and could be used to get the whole effort dismissed. In August 2019, the president and director general of IUCN wrote a firm statement of support for the assessment and its authors, but this did not end the controversy. In October 2019, several organizations that were members of IUCN sent a letter to the president, saying, “We are of the strong view that this report should not be regarded as a sufficient basis for the development of IUCN policy recommendations.” Those writing the letter raised two particular complaints. First, they said that several sections of the report seemed “to be biased toward the interests of those who intend to apply the respective technologies; some of whom were invited to be co-authors.” Second, they argued that the report avoided the most fundamental question, which was whether genetic engineering should be used at all in nature conservation. Instead, by talking about the introduction of GMOs and gene-drive organisms into wild populations without what they considered to be proper consideration of their implications, they suggested that the report effectively promoted their use. The letter’s signatories dismissed the report, even though it met IUCN’s terms of reference in offering an evidence-based review of both concerns and possible benefits. They argued that organisms transformed by synthetic biology should not be introduced into natural populations. Such actions would “entail the genetic engineering of the ‘germ line’ of bio­ diversity with the risk of disrupting the functioning of existing ecosystems and their future evolutionary dynamics.” They listed a series of impacts, including on animal, plant, and human health and biodiversity of value to Indigenous peoples and other local communities.10 This response to the IUCN report shows how strongly some opponents of genetic editing hold the view that organizations favoring synthetic biology are only engaging in conservation applications in the hope of winning a social license for synthetic biology more generally (with all their market-friendly nature-unfriendly products and processes). They believe that corporate support for conservation applications is designed to familiarize people with genetic technologies, in the hope of encouraging public acceptance and defusing any possible future opposition to possible commercial applications. One critique of genetically engineered trees, for

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example, portrays conservation’s dalliance with synthetic biology as a sort of “gateway drug” that will lead inevitably to less-nature-friendly applications. A critical comment on the IUCN report took as its title “Driving Under the Influence,” implying irresponsibility and lack of control in those doing the science, if not necessarily of those doing the review.11 Questions about potential impacts of synthetic biology on conservation reach a public already deeply affected by the long, bitter, and unresolved controversy over genetically modified (GM) crops and other genetically modified organisms (GMOs) in the 1990s. This was the first public exposure to genetic technologies, and it continues to shape public opinion about synthetic biology in many countries. In Europe especially, environmentalist protest led to widespread public opposition and tight regulation of GMOs. Many scientists interpreted the conflict as a victory for anti-science environmentalists, who deceived an ignorant public to whom the benefits of GM had not been properly explained. They saw the debate as a competition between irrational beliefs or “myths” and “facts.”12 But many other factors were also involved. GM crops were trialed without much explanation or public consultation and without consideration of the consumer or public perceptions or wishes. Many of the trials were run by private biotechnology companies who had developed (and who stood to benefit from) the crop varieties being tested. Furthermore, the research questions that the experiments were designed to address were relatively narrow: they did not, for example, pay much attention to the likelihood of gene movement from GM plants to nearby crops or wild relatives. Whatever the faults or limitations of the GMO testing and approval processes in different countries (which lie beyond the scope of this book), the fierce debate left a deep legacy of public distrust of genetic engineering, and an answering distrust of environmentalists and concern on the party of many scientists that the public was too gullible.13 In her book The Ethics of Invention, science and technology scholar Sheila Jasanoff makes a number of hard-hitting points about technology that speak directly to debates about synthetic biology and nature. First, she argues, “In all of its guises, actual or aspirational, technology functions as an instrument of governance,” changing human identity and relationships. Second, she highlights what she sees as the flawed idea that



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technology is politically neutral and outside the scope of democratic oversight. Lastly, she debunks the idea that technology “once invented, possesses an unstoppable momentum, reshaping society to fit its insatiable demands.”14 It is, as Jasanoff shows, impossible to separate technology from the social and political context within which it is developed and wielded. And it is essential that discussions about the governance of technology are made part of the democratic process. These findings set the stage for debates about how decisions should be made concerning the development, testing, and deployment of genetically engineered organisms, both in terms of their ecological impact and their use within conservation itself.15 There is an argument that the value of synthetic biology both to human welfare and to the economy of countries like the United States or China is so great that regulation should be led by the biotechnology industry itself. The main argument in favor of this is that it is in these companies that the real cutting-edge expertise in synthetic biology is concentrated. Technology is moving fast, and government investments in its science capacity have shrunk in many countries, as neoliberal styles of “downsized” government have advanced. Over the past three decades, the burden of regulatory science has tended to pass from government to the commercial sector, in systems of self-regulation in many sectors. Those proposing to introduce new technologies and products are expected to provide the scientific evidence on which decisions about their approval can be made. However, while this approach might work for established sectors dominated by blue-chip companies, the free-for-all world of synthetic biology innovation is clearly unsuited to such an approach. Letting innovation rip, in the tradition of garage computing, to see what new Microsoft or Google arises from the resulting storm of investment, may be a great way to build the biotechnology sector, but it does not offer much guarantee of a thoughtful assessment of the risks of genetically engineered species, or a sensible response to public concerns about them. At the end of the day, most independent commentators seem to believe that synthetic biology should be regulated from outside the industry itself, by governments. Scientists, environmentalists, industrialists, the media, and the general public agree on the need for a level playing field, a transparent regulatory regime, and clear mechanisms for public involvement,

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all with the aim of promoting a safe and sustainable world—and (if conservationists are allowed a voice) a biodiverse one. However, there are significant problems in asking individual governments to regulate technologies that are advancing so fast in so many countries and that promise bullish economic growth in the biotechnology sector. It is easy to call for international agreements on genome editing, or the release of edited organisms into the wild, but who should do such international regulation? The rise of national geopolitical interest, restrictive trade agreements, and national economic concerns make an appeal to internationally set rules increasingly unrealistic. In a world in which globalized corporations reach effortlessly across the globe, exchanging ideas, technologies, people, materials, and finished products from one side of the earth to the other, how can any organization hope to control the deployment of engineered biologies and organisms? The rapid arrival of synthetic biology on the policy scene, its shape-­ shifting nature, and the rapid evolution of its technologies make it particularly hard to fold into existing policy frameworks. There is as yet no agreed-upon international way of governing synthetic biology. That said, a number of international legal frameworks do exist that bear on this issue. Chief among these is the UN Convention on Biological Diversity (CBD) and its relevant “subtreaties,” also called protocols (with names that reflect where they were negotiated): the Cartagena Protocol on Biosafety, the Nagoya-Kuala Lumpur Supplementary Protocol on Liability and Redress, the Nagoya Protocol on Access to Genetic Resources, and the Fair and Equitable Sharing of Benefits. Other relevant treaties include the International Treaty for Plant Genetic Resources for Food and Agriculture, the UN Convention on International Trade in Endangered Species, and the UN Convention on the Law of the Seas.16 The Convention on Biological Diversity (CBD) has been discussing the need for international regulations for the use of synthetic biology, or even a moratorium on research, since 2010. Synthetic biology and especially gene drives were debated at the conferences of the parties to the convention in 2016 and 2018, both on the main floor and at many side events. Discussions were long, involved, and sometimes contentious. Arguments particularly concerned two issues. The first was the technology’s implications for the sovereign rights of countries to their own genetic



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resources (which are enshrined in the convention) and on the fair sharing of any benefits that derive from them—who owns the genetic information being used to develop synthetic biology, and who profits from its development. The other was the issue of biocontainment—the extent to which the proliferation of engineered of genetic modifications could be prevented, and who would be responsible in the event of such escapes.17 Years of debate and negotiation, inside and outside CBD working groups, have not yet reached any conclusion; indeed they have not even resulted in agreement on the definition of “synthetic biology,” because the definition selected would determine which of the existing treaties, accords, and regulations were brought to bear on the question (Should synthetic biology be considered as a new technology and a possible threat under the Cartagena Protocol, or simply as an issue of equitable benefits under the Nagoya Protocol?), and different interest groups have fought to postpone a decision on the definition for fear it would disadvantage their cause. The international community’s attempts to provide a framework for synthetic biology are at present effectively stalled. Yet it remains vitally important. Neither global biotechnology investment nor genes respect national boundaries, and many of the problems that synthetic biology is being used to address are global. International regulation may be difficult, but it is vital. The rise of synthetic biology and the abilities it provides to read and write DNA are putting strains on existing international agreements. For example, the structure of “access and benefit sharing” in international law, designed to support national efforts to sustain biodiversity, is based on the existence of a physical specimen. But the advent of digital DNA sequencing means that no physical specimen is needed once the genetic code of a species has been read. This separation of a specimen from its genetic code was not foreseen in the CBD and is causing urgent debate whenever the governments who have signed it meet.18 Given the challenges to international agreement, the weight of responsibility for the regulation of genetic technologies therefore still lies with national governments. The global political system is based on state sovereignty. States control natural resources in their territory and have responsibility for things that happen within their jurisdiction. Decisions about governance of synthetic biology inevitably, therefore, largely take place

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within this context, except where a common position is negotiated between states, as in the European Union.19 Across the world, national governments have approached the regulation of synthetic biology in different ways. The regulation of synthetic biology in the United States and the European Union exemplifies these differences. In both cases, thinking and legal frameworks have built on experience with previous innovations, particularly with genetically modified organisms. However, these previous frameworks are straining under the nature of the new synthetic biology technologies and products. Regulation in the United States, which emerged with the first GMOs in the 1980s, is complex. The Food and Drug Administration is charged with deciding if genetically engineered food, drug, and biological products are safe. The Environmental Protection Agency regulates the safety of genetic engineering applications in pesticides and microorganisms under the Federal Insecticide, Fungicide, and Rodenticide Act and the Toxic Substances Control Act. The US Department of Agriculture (USDA) regulates genetically engineered plants under the Plant Protection Act. Most GM crops in the 1980s and 1990s contained transgenes (genes from another species), and the USDA duly did its best to review those products. However, in 2016, it was asked to judge the safety of a white button mushroom that had been CRISPR-edited to slow the rate at which it turned brown. It declined to regulate, saying it only had responsibility for products containing transgenes, and the mushroom only contained genetic material that came from its own genome, although the DNA had been edited. The Obama administration commissioned a study of the problem by the National Academies, and the matter was left to the Trump administration, which by 2019 had only indicated that the “bio­economy” was a science and technology priority of the administration, without saying what it would decide.20 Countries in the European Union have taken a more restrictive stance on GMOs. A directive in 2001 regulated the deliberate release of GMOs, and another in 2009 regulated the contained use of genetically modified microorganisms, although some countries have extended it to all GMOs. The European Food Safety Agency evaluates the safety of new GMO products in terms of potential impact on human health, animal health, and the environment before allowing them to be brought to market. The assess-



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ment is based on scientific information presented by applicants, and it focuses on the molecular structure of newly created proteins, their functioning and potential interactions, the comparison of the GM plant with its conventional counterpart (phenotypic characteristics such as height and color and agronomic characteristics such as yield and nutritional values), plus an evaluation of potential toxicity and allergenicity and environmental impact. Once a GMO has been authorized, it normally receives a tenyear license for the EU market before reassessment.21 Since 2011, the European Food Safety Agency has started to assess organisms engineered using gene-editing techniques along similar lines. It is a painfully slow and painstaking process that presents a daunting challenge to biotechnology companies wanting to sell their products. Change has been proposed, but in 2018, the Court of Justice of the European Union ruled that synthetic biology should be regulated in the same way as GM. This decision was greeted with dismay by many in the research and agricultural communities who make the case that this will result in the stalling of synthetic biology innovations.22 Opponents of the Court of Justice opinion would prefer the European Union to adopt “product-” or “trait-” based legislation more like that of the United States. The British Royal Society has a long-standing position that the regulation of plant and animal applications for genetic technologies should be based on “the trait that has been introduced rather than on the process used to introduce the trait.” This difference between “product” and “process” is critical in distinguishing different approaches to legislation. The EU legislation was established more than twenty-five years ago based on the then-clear distinction between conventional and transgenic crops. New technologies blur such distinctions. Argentina, Canada, the Philippines, and Bangladesh have adopted product-based approaches, whereas Brazil, India, China, Bolivia, Australia, and New Zealand join the European Union in process-based legislation. In sub-Saharan Africa, biosafety regulatory policies have increasingly been aligned with progressive agricultural and rural development policies, with biosafety reviews placing greater emphasis on anticipated benefits rather than risks. There have, for example, been cases of expedited approval of GM crops for confined field trials or general environmental release.23 Regulation of the use and release of genetically engineered organisms

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is essential to avoid unforeseen and damaging environmental impacts and to ensure maximum social welfare benefits from novel genetic technologies. The thrust of innovation in synthetic biology is from private sector investment, but responsibility for assessing the environmental impacts of proposed innovations and monitoring their short-term and long-term effects must lie with governments, working individually and together through international agreements and forums. Inevitably, too, it is governments that will have to pay the lion’s share of any remedial actions needed if innovations have unexpected effects. To do their task of regulation effectively, governments need teeth, and for that they need science. Regulators need independent research capacity that enables them to understand and predict impacts and to monitor the outcome of their decisions. Risk assessments require modeling capacity and, above all, data. Baseline surveys and ecological and biological surveillance may be unfashionable areas of science funding, but they are fundamental to good decisions about novel genetic technologies and the engineered organisms they produce.24 The kind of science required may also need to change. Different areas of knowledge need to be integrated: wise decisions about engineered organisms demand the involvement of environmental scientists who understand their ecological context, but also the involvement of synthetic biologists who understand what genetic changes are involved, as well as researchers in the social sciences and humanities who understand how society and individuals will respond to them. Different parts of government should be joined up, to ensure that gaps in scientific knowledge are identified and that scientific findings are acted upon in an effective and timely way. These are not small challenges, even for wealthy industrialized countries, particularly in an era of austerity and restricted government expenditure. Developing countries need access to the same science and regulatory capacity, making the effective regulation of synthetic biology a challenge that requires unprecedented levels of international cooperation and support.25 Whichever organizations regulate the release of genetically engineered organisms, there are challenging questions about the risks associated with different kinds of applications. There is, for example, an issue as to whether



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genome modifications that are designed to spread, like engineered gene drives, should be regulated in different ways than less intentionally aggressive devices. An important dimension of this is the idea that their unwanted spread can be limited. As we have seen, this concern has led to the development of new kinds of engineered gene drives with limited capacity to spread or endure, such as “daisy chain” drives (which are self-­ exhausting), or “reversal drives” (which overwrite the original drive and replace the genes it carries). However, concerns remain with the concept of gene drives. Off-target, or undesired, changes continue to be a significant concern both for advocates and opponents.26 Another key question is whether all products of synthetic biology should be dealt with in the same way. The philanthropist Bill Gates has argued, in a paper with the optimistic title “Gene Editing for Good,” that “gene-edited organisms are not transgenic.” His argument, widely used by others, is that the basis of GM involved the rather crude insertion of genetic material from outside the organism (potentially from a species in a quite different taxon), while gene editing just edits existing DNA. This would suggest that such “cisgenic” editing (using genes and regulatory elements that already exist within a species) only makes changes that could in theory be introduced through conventional breeding, and for this reason should be regulated differently from edits that introduce exogenous genes or traits. This argument would suggest that gene editing and synthetic biology should not be regulated as a form of GM.27 Many critics of synthetic biology take a very different view, arguing that gene editing and genetic modification are essentially the same, being simply different ways to achieve the same genetic engineering ends. (Indeed, a critique published in 2012 by the ETC Group, Principles for the Oversight of Synthetic Biology, called genome editing “extreme genetic engineering.”) This argument suggests that GM and genetic editing should be regulated in the same way, because, given the level of genome analysis now available, and the potential to “edit” by base pair, it makes no difference whether new DNA sequences come from another organism (transgenesis), or by editing existing sequences to resemble those of another organism (cisgenesis), or are wholly new, designed by a researcher. Synthetic biology allows all genomes—as well as de novo genetic sequences— to be treated like machine code.28

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Many opponents of gene editing argue that all gene-edited organisms pose sufficiently serious risks to nature that they should not be released, even to conduct research on their impacts. Unsurprisingly, this notion of a moratorium on genome editing, even for research purposes, is rejected by most science organizations as preventing not only basic and applied research, but also research designed to understand the impacts and risks of wild release of genetically engineered organisms. As we have discussed, this position is also rendered somewhat moot by the large-scale existing agricultural deployment of gene-edited organisms.29 One of the main areas of difference between advocates and opponents of genetic engineering with respect to the release of genetically engineered organisms turns on the concept of the “precautionary principle,” or “precautionary approach,” and the way this should be interpreted. The idea of precaution has been a staple of environmental regulation since its introduction in the German Clean Air Act of 1974. The precautionary principle has been used in many international agreements, in a range of different formulations. The 1992 Rio Declaration on Environment and Development stated, “In order to protect the environment, the precautionary approach shall be widely applied by States according to their capabilities. Where there are threats of serious or irreversible damage, lack of full scientific certainty shall not be used as a reason for postponing cost-effective measures to prevent environmental degradation.” The European Commission adopted the precautionary principle in 2000. References to it are often used to justify a moratorium on genetic technologies, although the use of the concept in this blanket way is also criticized for unreasonably restricting potentially valuable innovation.30 Leaving aside the complex debates about the different ways in which the precautionary principle has been applied, there is no doubt that the idea that societies should be cautious about novel technologies is entirely sound. Synthetic biology offers new technological possibilities that seem to be powering an industrial transformation, with all that promises in terms of economic growth and commercial profit. In many sectors (most prominently food and medicine, perhaps also biodiversity conservation), these technologies offer net social benefits. However, they also bring undoubted risks, particularly (from a conservation perspective) to ecosystems and wild species. Precaution about these technologies is entirely justified.



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Speed may drive commercial priorities, but it exposes society and nature to risk. Precaution is needed to give time to learn about new technologies and build a shared understanding between scientists and the public, government, and industry.31 However carefully governments do their science and make regulatory decisions, they also need to take account of acceptability of new genetic technologies to the public they serve. A vital role for national governments is to balance the interests of powerful players, such as the biotechnology or agricultural industry, with the interests of citizens and smaller established businesses, including smallholder farmers. Many commentators suggest that decisions about the release of genetically engineered organisms, or the use of gene drives, should be made locally, by those stakeholders who will be affected, rather than be imposed by remote national or international bodies (or by corporations). Lay and local knowledge can also provide important evidence for making decisions. Community consultation has been an important element in development of several public health applications of synthetic biology, for example, mosquito control against dengue and malaria in Australia, and the genetic engineering of mice to decrease the transmission of Lyme disease to ticks and on to humans. Full consultation is important in winning public support for such interventions.32 There is a particular need for the regulation of genetically engineered organisms to respect the rights and interests of Indigenous peoples. Several indigenous groups have developed formal statements and declarations on genetic technologies. The Maori of Aotearoa (New Zealand), for example, have engaged in a thorough exercise to articulate their perspectives on genetic modification and how their cultural values might be used to analyze the risks and benefits. The escape from pens of genetically engineered salmon into the waters of Puget Sound led the Yurok Tribal Council of the Klamath River in the United States to declare its formal opposition to the approval of the production, sale, or consumption of all genetically engineered salmon.33 It is easy to suggest that all those who are interested in or potentially affected by synthetic biology products (the “stakeholders”) should be involved in decisions about their deployment. However, making decisions

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in this way is a complicated and sometimes fraught process. Not surprisingly, different stakeholders often hold distinct views. Individual scientists will hold their own views about the need for a particular application or its safety, and those working for commercial biotechnology enterprises or research institutes may well hold radically different opinions from those working for government or for the many environmental organizations that lobby against synthetic biology. Meanwhile, members of the public may have a measure of distrust of all scientists and bring quite different knowledge or sets of values into their views about novel technologies. At the same time, research shows that public attitudes to technology are not fixed. People can reframe their views in the light of their understanding of technological developments and the implications and potential of these developments for good or harm. This suggests that public consultation should be an iterative process. It is not something that can necessarily be done quickly, and certainly not hastily, at the last minute before a technology is approved.34 The way people respond to scientific information that is given to them is shaped by many factors, including existing knowledge, previous experiences, and perceptions of possible benefit and harm. Experiments on science communication show that basic outlooks and shared values are very important. In fact, positions on risk and reward can be less about critical appraisal than what Dan Kahan of Yale and his colleagues call “badges of membership in and loyalty to cultural groups.” So, even if unbiased information were available, its influence on decisions would be affected by the affiliations of those both providing and receiving the information. This makes good decisions even more complicated.35 It used to be taken for granted that scientists should and could provide objective factual information that would educate and inform the public and policy makers. However, psychologists have shown that rather than neutrally receiving information, people privilege what fits their preexisting worldviews, the social and political environments in which they live, and the beliefs or values they share with others. Increasingly, people live in a world of “fake news,” in which facts are often contested, values are disputed, and “experts” are disparaged. In the case of synthetic biology, it turns out that (at least in the United States) public attitudes are ambiv-



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alent and trust in science is not to be taken for granted. This makes it essential that consultation is broad, and shows both understanding of and respect for multiple perspectives and types of evidence.36 In 2018, the UK Royal Society carried out an extensive “public dialogue” process to assess public attitudes to genetic technologies. They found that the way people think about gene editing depends on what it was being used for. Over 80 percent of the respondents approved of genome editing to address otherwise incurable life-threatening conditions, such as muscular dystrophy; 69 percent approved of genome editing in plants to produce cheaper medicines; 70 percent supported making crops more nutritious as a way of supplementing poor diets; and 77 percent supported prevention of crop damage, for example, by fungal diseases. By contrast, only 28 percent agreed with using genetic editing to make cosmetic changes to animals (for example, micro-pig pets or fluorescent fish); and only 23 percent agreed with genome editing to create perfect fruit and vegetables. The Royal Society also discovered that public trust depends on who employs those scientists. People trusted university researchers the most because they were thought to be the most independent. Only 16 percent thought that scientists working within business organizations (or funded by businesses) were trustworthy.37 Public surveys also report mixed views about the particular issue of gene editing in wildlife. A 2019 survey in the United States showed that on average people perceived more risk than benefit in such applications, with a large majority thinking that gene-editing wildlife could be easily used for the wrong purposes. Those who strongly believed in the authority of scientific knowledge had more favorable views on the benefits and moral acceptability of gene-editing wildlife. Participants in this poll who opposed genetic engineering believed that such interventions “mess” with nature and allow humans to “play God.”38 This concern about scientists “playing God” is a common response to modern biotechnology—either made explicitly or implicitly, underneath other concerns. It surfaces in many other debates, for example, on birth control pills, transplantation medicine, or stem cell research. It seems to be an expression of deep concern about potential human transgressions of some kind of fixed limits associated with a certain order (which may or

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may not reflect particular religious beliefs). It speaks of the dangers of human hubris and the risks of unproven technology in inherently fallible human hands. To some, it questions the very definition of “life” itself.39 Such concerns often coalesce around issues of biosafety and biosecurity. Biosafety concerns potential damaging effects for workers and the environment due to accidental interactions with harmful synthetic biology products. Biosecurity on the other hand refers to risk from the potential deliberate misuse of synthetic biology for bioterrorism or biowarfare. Bio­ security issues tend to come under the purview of national defense agencies, which are increasingly concerned at the possibility that the new genetic technologies might be used to create biological weapons, in the form of new disease organisms or organisms with novel properties (for example, rapid person-to-person transmission). Gene editing no longer demands huge investment in laboratories, making it conceivable that people with relatively limited knowledge of the methods could modify (and “weaponize,” in the jargon) the genomes of disease organisms. Both concerns are a manifestation of the deep worries that by engineering wild species, scientists have the potential to cause tremendous and potentially irreparable harm.40 The problem both for scientists and the public, in assessing any of the possible impacts of genetically engineered organisms, is really one of uncertainty (not knowing what impacts will arise) rather than risk (the probability of outcomes that can be predicted). This distinction was made by Brian Wynne in his pathbreaking reflection on the fallout of radioactive cesium 137 on the British Lake District following the Chernobyl disaster in Ukraine in 1986. As the fallout cloud moved north and then west, radioactivity was deposited in rainfall on both the Scandinavian and British mountains. Radioactive cesium polluted soil and vegetation and accumulated in the bodies of Arctic reindeer and British Lake District sheep and in their meat and wool. The regulatory question (and the associated public debate) concerned what sort of threat this offered to public health and how to assess the risks. Wynne pointed out how different concerns got tangled together in debates about potential public harm from radioactive sheep. The concept of risk would have been relevant had the odds and the range of outcomes been known or accurately predicted. But this was impossible, because there was no previous experiment with contamination



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of this kind on this scale. The concept of uncertainty was more applicable because the probabilities of harm were unknown, even though it was possible to know what sorts of things might happen. Wynne also pointed out that both risk and uncertainty were different from ignorance (not knowing what we don’t know), which is inevitably a challenge with truly novel technologies. The novelty of genetic engineering suggests that society (scientists included) may not have the tools needed to be able to make good decisions.41 Unfortunately, people are not good at calculating and weighing different types of risk and uncertainty. As Douglas and Wildavsky point out in their book Risk and Culture, sense of risk is socially constructed, and assessments reflect the social and organizational groups of which people are part. Each side in a given debate is likely to think the other is biased or serving the interests of their favored organizations. When an issue like the application of synthetic biology to wild species is so deeply divided, the chance of quick resolution is remote. The level of disagreement currently being aired at international meetings certainly suggests that no quick agreement on rules is likely.42 The question of reengineering the genomes of wild species clearly needs a great deal of care. In this context, “wild” species need to be understood to include not only those of conservation concern but also less loved species such as mosquitoes, zebra mussels, or rats. Even microbes need to be understood as wild species, whether in a microbiome or not, for few might truly be considered domesticated (the yeasts used in beer or bread making being key exceptions). Decisions about conservation applications need to be guided by questions concerning efficiency, cost, and benefit (how well technologies might work compared to other strategies); risk and uncertainty (and therefore whether it is responsible to use them); and public acceptability.43 Decisions will need to be made carefully on a case-by-case basis. Genome engineering of wild species may be appropriate in certain places and times, but not wise or necessary in others. There might, for example, be a stronger case for gene editing in species (such as corals) that provide physical structure in an ecosystem or that are essential to ecosystem function. Others have suggested that the IUCN threat status of a species

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might be relevant to the type of genetic interventions that could be considered.44 Certainly, the same principles should be applied to synthetic biology as to other kinds of conservation action. There must be a strong sciencebased understanding of the conservation problem, the proposed solution, and its ecological consequences; there must be a broadly consultative and transparent prioritization process weighing the risks and benefits of genetic intervention; and there must be careful monitoring of effects and processes to ensure that management is adjusted in the light of them. It is hard to know how risks balance against any conservation benefits in advance. Before any release of an engineered wild species, there must therefore be rigorous testing, first in the laboratory and then in a contained natural setting.45 An example of the degree of care being proposed, and needed, is a 2019 study by the US National Academies on strategies to conserve coral reefs. They analyzed the risks, benefits, and feasibilities of different approaches, including genetic engineering. They recommended a flexible decision framework that reflected local ecological conditions and took into account inputs from a broad set of stakeholders. They suggested that deployment of any tool should be monitored and that management be changed if necessary. Perhaps their most valuable observation was that all tools should be considered and compared, so that the costs, benefits, and risks of synthetic biology applications are explicitly compared with other strategies.46 The acrimonious disagreements about the application of synthetic biology to conservation that currently dominate debate reflect concerns that lie much deeper than simply their impact, important though this is. Scientists do need to develop more reliable genome-editing techniques and safer gene drives and to undertake more extensive assessment of the risks of gene flow out of target organisms or ecological impacts. But if they did all these things, doing so would not be likely to result in immediate acceptance of their proposals. Public acceptance does not depend solely on better and better-publicized science. Daniel Sarewitz titled one of his articles “Science Can’t Solve It”—and that describes just where the world is in the debate about synthetic biology and conservation.47 In her book Can Science Make Sense of Life? Sheila Jasanoff writes that scientists working on genetics have used the metaphor of “the human



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genome as the book of life,” written in the “four-letter code of DNA,” thereby claiming for themselves the right to interpret its mysteries. Yet she argues that DNA represents only one of the books that inform human life, with at least four “other volumes,” those on “law, religion, political theory, and moral philosophy.” In our experience, many people have never heard about synthetic biology or its human medical and agricultural applications, let alone its potential uses for conservation. When told about the ways in which gene editing could be applied to conservation, they draw their responses not from science, but from their thoughts, beliefs, values, and experiences, all written in the languages of law, religion, politics, and philosophy.48 It is hard to know how broad the objections or endorsements for genetic technologies used for conservation purposes are. As we have seen, most people do not yet know anything about these technologies or their impacts on ecosystems and species and their conservation. Yet these technologies are currently subject to very intense international discussion and scrutiny. It is hard to think of any other conservation tool that has received such scrutiny. Hopefully the global attention will bring with it greater understanding of the science and its potential for the harm or good of the natural world. The idea of genetically engineered wild organisms modified not for human profit but for the “good of nature” opens up a new ethical landscape for conservation. Such modified organisms will be what the cultural theorist Arjun Appadurai called “things” with social lives that are tightly wrapped into human culture and society. They will create, in their turn, novel social and cultural practices and norms. Decisions about such interventions in the natural world will quite rightly be made through debate by communities, Indigenous peoples, national governments, and global bodies. If such debate is permitted and informed by clear arguments and strong evidence of diverse kinds, there is some hope that the decisions reached will be careful, widely accepted, and have beneficial outcomes in terms of the well-being of human society and nature.49 But the questions raised by genome editing go beyond the important practicalities of regulation. Genetically engineered life not only has the power to transform economies but also to change the way people think about nature, that ever-malleable concept. Made and released, genetically

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engineered creatures would become part of ecosystems. They would be living, evolving, ecological actors playing out their roles as part of a nature that evolved without them in their current form. Beyond the important question of their ecological impact, there lies the issue of what kind of nature engineered organisms really are. That question reaches deep into the heart of conservation and into humanity’s idea of what it means to say something is “natural.”

9 Conserving Strange Natures

At the Ballard Locks Fish Ladder in Seattle, Washington (United States), you can sit underground on a clammy concrete bench and look directly through a thick plate of glass into the murky green-tinged water moving from Lake Washington down to Puget Sound. In June or July, sockeye salmon speed past, or pause as if to catch breath, before going on their way upstream. They are returning from feeding in the ocean to their spawning grounds in the Cedar River Watershed. The Ballard Locks, built by the US Army Corps of Engineers, were closed in 1912, disrupting the salmon run in the Duwamish River. The fish ladder, opened in 1917, consists of a series of pools between which the salmon can move, giving them a way around the locks. In the first week of July 2019, over thirty-five hundred salmon of three species (sockeye, coho, and chinook) maneuvered around the locks, the busiest in the United States, and swam on toward their mating destination in the gravel beds of streams fed from the Cascade Mountains.1 Salmon are born from eggs laid in the gravel of streams or lakes and grow into free-swimming fry. Though some stay close to where they hatched, the majority move to other freshwater environments or the ocean. After a period of feeding and growing, adults follow the scent of the stream where they were born and run up it, to spawn in their turn. That

Sockeye salmon 191

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return is an innate response, coded into the salmon’s DNA. Every year, the cycle repeats. Unless it doesn’t. It was not until the advent of scientific programs to mark salmon by clipping their fins or attaching threads or tags, that researchers could track individual fish movements. They soon began to notice that not all individuals returned to the river they had come from. Some appeared to make mistakes, returning to the “wrong” place. The proportion doing this is small, ranging from about 10 to 30 percent depending on species and population. This behavior, called “straying” is not a result of faulty DNA or, in fact, a mistake at all, but is an extraordinarily important part of the evolutionary success of salmon.2 Much of what is now salmon habitat was under glaciers over ten thousand years ago. As the ice melted, rivers started to create prime salmon habitat in new locations. This could only be used if fish from an existing spawning population went the “wrong” way to check out a new place. And it is to their genetically coded predisposition to seek novel waters that today’s extraordinary—and in many cases unfortunately now imperiled— salmon runs owe their existence. Straying is not just behavior relevant to the remote past; it is still at play. Its power is demonstrated by the return of salmon to the north fork of the Toutle River after the eruption of Mount Saint Helens in Oregon on May 18, 1980, which boiled any salmon and eggs in the river. Salmon also returned to the Elwha River on the Olympic Peninsula in Washington State after the removal of the thirty-three-meter-tall Elwha and Glines Canyon Dams in 2012 and 2014 by the US National Park Service. In Alaska, salmon are now colonizing streams newly opened as glaciers retreat, recapitulating the postglacial pattern of colonization. In a less desirable context, straying behavior has allowed escaped farmed Atlantic salmon to invade and colonize some of southern Chile’s rivers—a continent away from their natural home. Salmon colonization demonstrates the adaptive capacity that is so vital to allowing salmon to continue to evolve to become new salmon pop­ ulations or ultimately perhaps new salmon species. Genetic variation— sometimes called “adaptive potential”—is a critical element in nature’s ability to respond to a changing world. Though the details are not always clear, the amount, distribution, and functional significance of genetic vari-



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ation can all contribute to the ability of a species to respond to environmental change.3 In Pacific salmon, as in other species, the diversity of the genome is fundamental to evolutionary potential. Not only do genes within different individuals reflect different local adaptations, but genomes also contain substantial lengths of “spare” or “junk” DNA. These do not appear to serve as “genes” (meaning they do not appear to code for any particular trait), but recently, as we have discussed, it has been shown that they are the genetic raw material from which genes can be made, a kind of “reservoir” for evolution.4 As we have seen, conservation geneticists have developed a growing suite of genomic approaches to conservation. The importance of diversity in genomes to the survival of threatened species has become abundantly clear, as is the fact that genomic interventions that decrease the potential for adaptation made possible by such diversity are likely to decrease the chances of survival. Active management techniques (such as captive breeding, genetic rescue, assisted colonization, or microbiome conservation) are either already used or are contemplated, to counteract human reductions of genetic diversity in threatened species.5 Techniques of genome editing open up another possibility. The conservation-genetics literature has so far had little to say about the possibility of actively altering the genomes either of threatened species or those that threaten them. But, as we have seen, scientists are asking if there is a case for active, intentional, genomic intervention to reshape the genomes of wild species and their microbiomes to improve their chances of survival. Organisms that conservationists value, unlike disease-causing microbes, are often unable to adapt to keep pace with human-driven environmental changes. Genetic engineering could perhaps speed up or direct their adaptation in response to the damage humans do to ecosystems. Arguably, the mess of the Anthropocene will not fix itself, no matter how assiduously other conservation approaches (for example, protected areas or trade restrictions) are pursued.6 Engineering the genomes of wild species, such as those declared as “pests,” seems to be here to stay, driven by the power of the agricultural and biotechnology industries, and, as we have argued, it urgently needs addressing by conservationists. It is scary if genetic engineers and entre-

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preneurs do not understand ecology or the implications of their work in the wider landscape. Conservationists therefore need to engage in debates about the release of genetically engineered organisms and contribute to wider understanding of the biological and social implications of synthetic biology. This we have already discussed. Here we look at the implications of such gene-editing technologies for conservation itself. It goes without saying that the techniques of gene editing are highly artificial. But then, as we have seen, artificiality has a valued and carefully considered place in conservation practice. We are, for example, familiar with the idea of habitat creation as a conservation strategy. In the East of England, the Royal Society for the Protection of Birds (RSPB) owns Lakenheath Fen, a nature reserve on the edge of the huge former wetland of the fens, all but a tiny fraction long converted to agriculture. Before the RSPB bought it in 1995, the three thousand hectares of land that comprise the reserve had been intensively farmed. They used heavy earthmoving machines to reshape the site for conservation, turning carrot fields into water bodies, reed beds, and wet grazing marshes. The result of all this work is a stunning bird reserve. One cool, cloudy early summer morning a few years ago, we were walking through the reserve, watching a small falcon flying high over a block of mature poplar trees, wondering if it was a hobby, a rare summer migrant to the United Kingdom. We were discussing naturalness and artificiality in conservation, a discussion that marked the beginnings of this book. We asked ourselves in what senses was Lakenheath Fen—with its bulldozed carrot fields, carefully moderated water tables, and created reedbeds— natural? For all the creative engineering needed to make the physical environment at Lakenheath Fen, the birds themselves came on their own. As well as hobbies, there are bitterns (a secretive heron whose UK population had fallen to just eleven birds in 1997) and marsh harriers (a rare reedbed bird of prey). The colonization of the new reserve was, at least in this sense, natural. One bird that arrived was the statuesque common crane. This had died out in the United Kingdom in the medieval period, although it was known as a rare visitor, when migrating birds overshot their mainland European breeding grounds. In 1979 a crane appeared in Norfolk, in eastern England, and in 1982 a pair bred there for the first time. Slowly their



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numbers grew, boosted by a conservation project in western England that raised crane chicks in a protected environment (just like the whooping cranes described in chapter 2). The first cranes were released into the wild in 2010. As if to offer a seal of approval to Lakenheath’s creative conservationists, cranes turned up to enjoy the new wetland, and in 2017 they nested. This particular event, too, was a natural colonization, but one made possible by a great deal of effort elsewhere in the country. The wild cranes at Lakenheath were probably raised on what was essentially a highly specialized bird farm. Their “wild” behavior in colonizing Lakenheath was the product of the cleverness of the techniques used to rear them and the availability of a former carrot field converted into a place where they could breed.7 Nature at Lakenheath is clearly a mix of naturalness and artificiality. The result is nature that is wild, but (as we put it in this chapter and the book) strange. Here, as in so many cases, conservation involves both the artificial assembly of natural parts and the natural evolution of artificial parts. The question of whether it is natural to use a bulldozer to create a reedbed to attract wild cranes to converted carrot fields may seem a long way from the application of gene editing in conservation. But it is not. Both raise questions about the place of artificial methods and modern technologies in shaping the nature of nature. Both call attention to the tension between novelty and naturalness. Genomic tools allow scientists to make deliberate, purposeful and intrusive incursions into the genomes of wild species. They make it possible to imagine changing genes to “save” species. So the key question for conservation must be what to think about using technologies—made things— in order to save nature. And if we are happy with bulldozers and planted reedbeds, in what ways are they different from genetic technologies and human-transformed organisms? Growing knowledge of the genome, and the accessibility of technologies to change it, are repeating and amplifying existing debates about naturalness in the conservation of ecosystems. There is a long history to conservation’s efforts to define and protect naturalness. This issue lies at the heart of conservation, helping shape the complex and diverse sets of values that conservationists hold. The power of genome editing makes the answers to those questions newly important. As ethicist Ronald Sandler observes, genetic engineering has the potential

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to restructure conservation ideas, practices, and value positions. Put simply, what is natural in a world of genome editing?8 In Greek myth, Prometheus was a Titan who stole fire from the Gods and brought it to humanity. For this gift of technology, he has been celebrated as the father of science and civilization. But for his boldness he endured a terrible punishment, condemned by Zeus to be chained to a rock to suffer but never die as an eagle ate his liver, only for it to regrow each night and be devoured again the next day. Much of what we are as modern humans has its roots in the command of fire. Yet fire and the many other technologies it symbolizes have often been abused. For this reason, Prometheus has also come to represent the ambivalent role that science and technology play in human society. Mary Shelley chose The Modern Prometheus as the subtitle for Frankenstein, and the novel highlights the unfolding moral choices and unforeseen ethical responsibilities that follow scientific advance.9 Uncertainty about technology lies behind the emergence of environmentalism as a global movement in the twentieth century. It is a central theme in many of the works that have inspired environmental concern through the decades, including Man and Nature, A Sand County Almanac, Silent Spring, The Limits to Growth, and An Inconvenient Truth. Ambivalence about technology has provided the script of a Greek chorus to accompany the last centuries of human progress, reminding us that automobiles, nuclear power, computers, artificial intelligence, and a thousand other technical innovations offer risks as well as benefits and bring the likelihood of unintended consequences. Nowhere is this ambivalence more relevant than in the potential application of synthetic biology to the conservation of wild species.10 There is no doubt that the idea of using synthetic biology tools to deliberately alter the genomes of wild species is a tricky one for conservationists. But is its novelty and weirdness a sufficient basis for rejecting such interventions? Genetic technologies are certainly novel, and their impacts are largely unknown, which is why society needs to be very cautious about their use. The pragmatic argument for the use of synthetic biology in conservation does not concern itself with whether technologies are natural or not. It focuses instead on rationales of efficiency (Do they enable



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conservationists to reach their goals more effectively than other interventions?) or necessity (Will conservation efforts fail without them?).11 One common argument about gene editing is that it is different in scale but not in kind to what humans have done before. After all, people have been altering the genomes of species for millennia, long before they even understood that something called a gene existed. Moreover, clearly the genomes of a number of “wild” species are already human-influenced, through adaptation to human-made and human-influenced environments (from cities to climate-changed “natural” habitats), and others are being affected as part of agricultural pest and disease-vector control programs. Yet there is a difference between selection pressures operating gradually within a complicated mix of social and ecological forces over decades, centuries, or millennia and direct genetic manipulation performed in a laboratory in years or months. Gene editing is not just faster, but it is also deliberate, planned, and targeted. The ways in which the evolution of diversity happens naturally through mutation, natural selection, and evolution, even when in the context of the Anthropocene biosphere, are different from the “artificial” gene hacking of synthetic biology and its intentional genetic alterations, let alone its potential to write entirely novel genetic code. The degree of human purpose in the shaping of the genomes of wild species matters in terms of the balance between naturalness and artificiality. There is a difference between the accidental impact of humans on the genomes of wild species (for example, in urban environments), and the deliberate manipulation of those genomes according to some premeditated human plan. The genomic changes being proposed and tested by the synthetic biology industry extend and accelerate the millennial record of domestication by humans to such an extent that arguments about continuity are threadbare. There is an obvious difference between the blind watchmaker of evolution and design by human editors. It seems clear that there is a question about whether a wholly novel synthetic organism can ever become “natural.” The critical point for conservation is subtler: Does this question about the “naturalness” of engineered organisms also apply to wild species whose genomes are manipulated? In particular, does it apply if genomes are edited for conservation purposes?

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In thinking about “naturalness,” the issues raised by the genetic engineering of wild species are different from those raised by the engineering of already domesticated species. The genetic tools are essentially the same, but the context and risks are very different. The genomes of relatively few wild species are known. Knowledge of the genetic traits that underpin ecological interactions between species is even less available. The effects of genome editing are therefore harder to predict than in a well-understood genome (for example, in the domesticated pig, or a laboratory mouse). Many wild species of conservation concern are rare with consequently eroded genetic diversity. The level of risk of experimental genetic interventions in such species is hard to know. The degree of threat that a species is under might suggest the desirability of a genome-editing strategy as a technique of last resort, but also the need for precaution. It is a reasonable conclusion that the genetic editing of wild species should be done with particular care, if at all.12 Gene-editing technologies may not be any more unnatural than other technologies regularly used in conservation, but arguably they are very different. So, for example, one might compare the use of CRISPR to edit the genome of black-footed ferrets to increase their resistance to disease with a chainsaw used to control invasive trees. Both the CRISPR module and the chainsaw are technologies. Both have important biological and ecological impacts when applied by their human handler. Yet they are very different in their effects and how easily those effects can be observed. The chainsaw can only cut one tree at a time, it takes a while to do it, and the results are immediately obvious to any observer. If someone thinks the cutting is unwise or illegal, there is the potential chance that they can influence the management decision. CRISPR is able to restructure a whole genome, but it does it in a laboratory, and it can only be observed by the scientist making the change. Moreover, its effects are not confined to one individual but can persist down the generations, as human-caused genomic changes undergo natural selection when the altered organism joins its ecological community. The two technologies trigger different concerns about risk and the willingness of society to accept unknown technologies.



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Scientists and opponents of genetic editing of wild species, and gene drives in particular, are both given to the language of hyperbole in describing technologies like gene drives. One news piece in the prestigious journal Nature opened an article on the use of gene drives in mosquito and malaria control by saying scientists had been “trying for eight years to hijack the mosquito genome. They wanted to bypass natural selection and plug in a gene that would mushroom through the population faster than a mutation handed down by the usual process of inheritance.” Another article described gene drives “clashing with evolution.” Such language suggests the idea that nature is on its own path until disrupted by CRISPR and other powerful genomic tools. It contrasts gene editing with natural processes, implying that the science of gene editing is unnatural. And this is exactly why many opponents of the technology are so opposed to it—its unnaturalness. Some proponents of the technology agree that it is unnatural but celebrate the powers it offers.13 The most fundamental question for conservationists is whether gene editing is an appropriate tool for them to use or not. Many thoughtful people answer this question with a clear “no.” This position reflects an essentialist view of nature, as something that exists outside of the human realm and whose value derives from that independence. Any application of technology is seen to break down the human-nature distinction, weakening the sense of nature as independent of human interests and threatening to erode its value. In her classic book Purity and Danger, Mary Douglas suggested that “dirt” be defined as “matter out of place.” A weed is often defined as a plant growing out of place. So how should we imagine a gene out of place? Douglas couples purity with danger—the danger that comes from despoiling purity. The concept of purity is a powerful one for humans and one with a long and complicated history in many contexts. Though conservationists mostly do not talk in terms of purity, it figures in our thinking in coded form. One can see it in the popular enthusiasm for “wilderness” and the need to believe that there is a place free of human influence, a place clean and pure. This trope has arisen time and time again in Western civilization from the search for the Garden of Eden onward.14 As we and many others have argued, wilderness is a cultural concept.

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Yet despite the overwhelming evidence documenting the depth and extent of human impact on nature, the notion of a pure and untouched nature waiting to be protected retains a powerful draw. The genomes of wild species can be thought of in the same way that people think about wilderness, as something that is pristine and little affected by human activities. To borrow the language of the US Wilderness Act of 1964, undomesticated species might be described as “untrammeled.” The architect of that legislation, Howard Zahniser, defined this term to mean not being “subjected to human controls and manipulations that hamper the free play of natural forces.” The major difference between wild and domesticated species would then lie in the extent of direct and purposeful human action involved in shaping their genomes. The genetics, physiology, movement, reproduction, and ecological interactions of wild species are still, in many instances, little affected by intentional human actions.15 Robert Elliott points out that the difference between the original and a perfect copy of a work of art lies in its genesis. By analogy, he suggests that the value of a natural landscape is its “causal continuity with the past.” He argues that human interventions, even those intended to protect or restore nature, inevitably reduce this value, a process he memorably called “faking nature.” If the value of nature lies in its origins, in the kinds of processes that brought it into being, an important reason to conserve an ecosystem or a landscape is the fact that it was not brought about through human intervention.16 The same argument can be made about genetic intervention in the genome of a wild species. Many writers have explored the ethical significance of nature’s independence of human decision or will. It is a widely accepted principle in conservation that ecological and evolutionary processes have value in themselves. Philosophical debates about such “intrinsic values” get complicated, but the important thing is that value is held to be associated with the independence of nature’s physical and biological processes and the species and ecosystems within which they occur (and which they create). To follow Elliott, the argument might go that if a wild organism has (mostly) evolved independently of human intervention in design, creation, and material composition, part of its value lies in this independence. This value is reduced by genome editing.17 By contrast with a wild species, a synthetic organism is an artefact,



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designed and engineered by humans. But all organisms whose genomes have been edited share a degree of this “artifactualness.” If independence from human intervention is an important source of value, then genetic engineering, with its human design and control, surely undermines that value. Moreover, it represents the attempt to forcibly adapt the species to the world humans have created, a very different thing from conservation’s ostensible aim of adapting human lifestyles, industries, and resource uses to ensure the survival of that species. An example of this argument is the suggestion that genetic adaptation to problems such as anthropogenic climate change compromises both the wildness of the species concerned and the places where they live.18 The idea that conservation must protect what is “natural” is understandable, but the distinction between what is natural and what is artificial no longer provides a sound guide to thinking about people and nonhuman life. A range of highly artificial acts, using fences, guns, traps, poisons, chainsaws, and many other artificial devices and techniques, are all routinely used in the name of conserving nature. Except for a very few places, almost all of them already in protected areas, the nature that survives on earth is subject to human management or influence. Those who care about nature have come to accept, however grudgingly, that most ecosystems and species are affected by human activities. Genomes are subject to influence in just the same way. What we know about contemporary evolution, hybridization, horizontal gene transfer, and the relative openness of species boundaries makes this quite clear. Animals or plants in zoos, gene banks, or seed banks are isolated from these changes and are effectively decoupled from the dynamism of the world from which they were removed. Those in the wild continue to evolve, responding to natural and human-influenced climate change and other features of the Anthropocene. The idea of the genome as analogous to wilderness is a powerful idea, but it fails us for the same reasons that it does at the landscape or ecosystem level. Conservation cannot be limited by an essentially vain search for the pristine and by the attempt to maintain that idealized state or restore nature to it.19 As we have seen, ecosystem restoration is an increasingly important

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element in conservation policy worldwide. Restored ecosystems are inevitably different in their species composition and function from any present or past system, with all the complexities that means for their conservation and management. As we have seen, more and more surviving biodiversity exists within novel ecosystems—hybrids combining the influence of nature and people—and in the human-influenced genomes of the species that comprise them.20 To say this is not, as some critics suggest, to abandon the ethical claims that nature has value independent of human society. It is to recognize the need to take the scale of human capture of natural processes seriously and to acknowledge the ethics of care. There are risks to seeing all of nature as a garden (even a rambunctious one), for the idea carries a vast freight of values with it, many of them narrowly anthropocentric. Human management of nature can be careful and creative, and the results can be beautiful. Humanity loves the versions of nature it has made for itself—parks, gardens, tree-lined streets, and landscaped rural estates. We know these places are artificial, and we don’t confuse them with wild places. Our love for them reflects the way they provide rich sights, sounds, tastes, and smells that are not human-made (even if they are human-assembled). These are environments that we value for their naturalness, even though we recognize the extent of human choice and craftsmanship in their creation and the degree of care invested in their management.21 We see examples of things that mix the human with the natural every day, in every field and forest and every city in the world. In an outside courtyard of the Honolulu airport, a small pond surrounded by trees is visited by wild ducks, paddling contentedly as nervous passengers search for their gates. In Australia, some people argue that the introduced Australian dingo should now be accepted as its own full species and therefore welcomed as a member of Australia’s native fauna. Late at night coyotes in US urban centers, and in the United Kingdom, red foxes, emerge from lairs in the interstices of urban infrastructure to forage among the discarded fast-food containers. Such environments are often unhealthy for wildlife, as they are for people (crows that feed on such human trash have even been shown to suffer from high cholesterol levels), but they thrive nonetheless. Natural-anthropogenic hybrids abound, increasingly the rule and not the exception.22



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As we have shown throughout this book, people have so thoroughly inserted themselves into the natural world that it is often difficult to tell where their influence stops and the “true” natural world begins. No part of “nature” should be imagined or managed as if it were utterly separate from human artifice (and if it were, the very act of management or designation would compromise its un-human-ness). Decisions about managing nature have to accommodate many complex combinations of the natural and the unnatural. Take the example of the suburban deer around big cities in eastern North America. Whitetailed deer are common in New York State, where Kent used to live. In a landscape of one- or two-acre house lots, the deer are a great nuisance because of their appetite for garden flowers. They are thought of as a wild nuisance: unwelcome nature in a valuable and managed space. But the mix of woodland, lawns, and house gardens in this landscape is ideal habitat for white-tailed deer, and as a result, deer reach exceptionally high densities—densities that are not “natural” at all. Moreover, because so many deer are living so close to people, many are killed on the road. In a weird act of human-natural nutrient recycling, New York State road crews pick up the dead deer and compost them, making the resulting mulch available for suburban residents to enrich their gardens, providing more food for the deer. Such human activities are causing ecological changes that force very uncomfortable reflections on how we imagine the natural world and on the adequacy of the fixed categories we like to divide it into. As a result, conservation is all the time being pulled in two conflicting directions. On one side is the pull of preservation. This is reflected in the long tradition of preserving nature, protecting it from the powerful forces of human use and their consequences (including, nowadays, climate change). The urge to preserve tends to be based on a sense of moral responsibility for those forms of life that share the planet and on strong aesthetic support for the experiential and existential values of wild animals and wild places. This side of conservation is animated by a strong desire to keep these things in their present state, or to restore ecosystems to some specified past state, before critical human impacts. The reality of past human-caused extinction and the specter of imminent future extinction are powerful motivating forces for this position.

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But conservation is also pulled in another direction, because the nature that people value is not a fixed entity. Species and ecosystems and the genomes that dictate their forms and compositions are constantly changing. They are “products” of evolution, but not in the normal sense of the word, as if evolution were a manufacturing process that turns out finished articles that humans can admire, use, preserve, or destroy. Biodiversity is not fixed. Species and ecosystems are way stations on a never-ending evolutionary journey. Even without human intervention, physical environments change, and with them the populations of different species and the composition of different ecosystems that they comprise. Evolution never stops. Species adapt, as natural selection acts on genetic variety. No one waypoint is final. The things people love—a buzzard rising over a Cambridgeshire woodland, or the alewife migrations that surge up the rivers of Maine—are transient. They will be here, if we are lucky, for scores of human lifetimes, but they are impermanent in evolutionary time. Synthetic biology raises specific issues relating to the naturalness or authenticity of species that have been reengineered at the genetic level. If scientists add the genes from wheat to American chestnuts and restore them to forests they once dominated, is this the restoration of real American chestnut trees or something different and less authentic? If things are hybrids of the made and the evolved, does this disqualify them from counting as “real” nature? And, to take a more extreme example, what if scientists coax cells to produce a novel organism that has not shared evolutionary past with any other living organism? Would the resulting living thing be an authentic part of nature? If it were released, should it be protected as a wild species? If not, how many generations of independent life would have to pass before it might win that accolade? And if the novel genome were part of a modified microbiome responsible for saving a species of endangered frog, would that make its acceptance easier? There is obviously a risk that the power to engineer the genomes of organisms will undermine conservation efforts based on the idea of “nature” separate from and threatened by human action. Genomes can be restructured in ways that cannot be seen except by using high-tech scientific methods. Both the public and conservationists might see genetically engineered “wild” species as less natural and less valuable—less deserving of protection—than those that have not been touched. The proliferation



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of such engineered species might result in a lack of clarity about what had and had not been changed in different species, potentially undermining the whole idea of the conservation of nature. So, to take an extreme example, if a re-created mammoth is really a genetic hybrid—an Asian elephant with some of its cells reprogrammed to match the mammoth genome—should we treat it as a wild species? Or would it simply be a hairy elephant that looked sufficiently like a mammoth to pass cursory inspection or to sell tickets at a theme park? The resulting animal, though possessing the configuration of the mammoth that was lost, will have been almost completely severed from the long evolutionary history of mammoths before they went extinct. It is possible that this human-created mammoth might perform the ecological functions of long-gone mammoths, but would that be any more authentic than programming a robotic bulldozer to push down woody vegetation, or a drone to deliver fertilizer? In addition, the de-extinct species would lack the original’s physiological, behavioral, and microbiome signature characteristics. Some of these might be restored, but not the whole thing.23 The determination of what is authentic or not, what is natural or not, and what is or is not ethically acceptable as part of conservation is far from straightforward. This is the tension that faces conservationists as they come to terms with the genetic-editing revolution. Their instinct to keep nature the way we think it should be clashes with the need to let it adapt. Should conservation try to lock down nature in the form it has taken in recent centuries, or should we open our hands to allow dynamism, change, and evolution? And if the latter, should we set limits to this openness? Nature is irredeemably strange in the era of genome editing. It is hard to know what it is and therefore what we should be doing about saving it. Into this world so marbled with human impact, knowledge of genes and genomes further confounds the choices for conservation. Genomes are not static any more than are ecosystems. They change constantly, forever kneaded by evolution, hybridization, horizontal gene transfer, epigenetics, and undoubtedly other processes that are not yet understood. In the era of genome editing, genetic diversity takes center stage in debates about nature’s future.

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At this time in the complicated and careless human history with the natural world, we need a new way to understand the interplay between nature and human intentions and actions. To know how to respond to gene editing, conservationists need to be able to think about this at the level of the genome. One idea that is helpful comes from ecosystem restoration, the idea of rewilding. As we have seen, rewilding projects now exist in a number of countries that are releasing wild carnivores or herbivores or domesticated grazing animals to try to re-create the ecological functions of “missing” species. Conventional ecological restoration tries to shape and manage ecosystems to resemble selected reference systems. The aim of ecosystem rewilding projects is to restore nature’s own dynamics (for example, geological processes of erosion and deposition and ecological processes of herbivory or predation). As we have seen, such an approach may make it difficult to predict exactly what pattern of species and ecosystems will emerge. The argument is that it may be enough to set the target of restarting processes like river flooding, predation, or grazing and allow the detail of outcomes to emerge over time. After all, a key dimension of “wildness” is the idea of land that is “self-willed,” developing in response to the interactions of its species as they adapt to each other and to the physical environment.24 Just like ecosystems, genomes are also dynamic and constantly changing in response to selection pressures and other processes. Human management of wild species can affect breeding, mortality, access to food and water, ranging patterns, and body size in animals and a variety of equivalent parameters in plants. Such changes could be said to decrease the wildness of a species—to the extent that its behaviors, ecology, and evolution are affected, directly and indirectly, by human practices. Releasing these human practices could be said to allow a species to become more natural and regain some of its self-will in ecological and evolutionary behaviors.25 Where conservationists manage ecosystems and species, they also manage genomes. As with ecosystem management or restoration, human genetic manipulation ranges from controlling natural processes to enabling them. One approach to genetic editing is to control the makeup of the genomes of wild species and the traits expressed to try to maintain them in what is thought to represent their “natural” state. Another would also seek to



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maintain genetic diversity, but with a different purpose. It would be to allow natural processes of selection and evolution to act upon the greatest diversity of evolved genetic patterns. By analogy with ecosystems, this could be thought of as rewilding the genome, working with natural dynamics and processes to sustain living diversity and the processes generating it.26 In a profoundly human-transformed world, a black-and-white distinction between “natural” and “artificial” at the level of the genome is no more satisfactory than it is at the level of the ecosystem. Our world is irredeemably gray, and with genomes (as with ecosystems) it is the “causal continuity with the past,” in terms of evolution, that matters.27 The diversity of the biosphere has evolved over geological time and continues to evolve today. Many human activities reduce diversity and also influence the direction of its evolution, but the core process of evolution continues, creating and changing life as the planet changes. The key objective for conservation is therefore to maintain the diversity and, through it, the evolutionary and ecological processes on which life depends, the link between the evolutionary past, the fraught present, and the uncertain future. The continued capacity of natural systems, from genes to ecosystems, to adapt and evolve lies at the heart of conservation’s concern for nonhuman life. So engineering the genome of a frog to increase its resistance to chytrid fungus will embed a novel or artificial element into its naturally evolved genetic portfolio. However, the frog would hopefully survive and continue to breed rather than go extinct. Almost all the rest of the genetic information encoded in its DNA, evolved over millions of years of evolution, would also still exist. As the frog continues to evolve, all that genetic material would go on generating variety on which natural selection could continue to act through myriad successive generations. The vast majority of evolution’s handiwork would persist, and the genome would be free to evolve into the future. The act of editing the genome of an endangered frog would release the natural processes of genetic variation, ecological competition, and natural selection. Like the release of a missing apex predator into an ecosystem, the edit would allow a cascade of biological processes to run. It might even restart a process of evolution that was slowing to a standstill as the chytrid fungus pushed the frog population toward zero.

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This way of thinking about rewilding the genome extends existing approaches to genetic rescue by seeking to secure the genetic basis of the evolution of responses to novel selective pressures. It supports the capacity of threatened taxa to respond both genotypically and phenotypically to environmental changes. It allows genomes to continue to evolve as part of the wider evolution of diversity, rather than constraining them to reach any specific human-defined outcome. Gene editing may therefore have an important role in enabling species to survive and ecosystems to self-organize in a world increasingly shaped by humanity. Such an evolving genome would be considered to be “self-willed” in the sense that it would not be subject to ongoing direct human intervention.28 If conservation were to follow this line of thinking, it might avoid the trap of seeing gene editing as yet another technology that can be used to tie nature down to human purposes, whether that is agriculture, biotechnology, or even the conservation of rare species. Synthetic biology might be used to support or assist evolution, sustaining evolved genetic diversity and enabling nonhuman life to evolve as independently of human influence as possible. The sun of early winter gilds the tall dead grass on the rolling South Dakota hills of Wind Cave National Park. In a sheltered swale, a group of female bison and their eight-month-old calves graze or rest in the warmth. The park, the seventh national park established in the United States, was created in 1903 to protect a deep cave system and a fine remnant of central North American prairie. President Teddy Roosevelt, who signed the bill creating the Wind Cave National Park, was part of a group of people who formed the American Bison Society in 1905 as part of the New York Zoological Society (now the Wildlife Conservation Society). The Bison Society’s purpose was to address the imminent extinction of American bison. Its numbers had fallen from thirty million to fifty million animals when Europeans arrived, to a few hundred by the end of the nineteenth century. The culprits were commercial slaughter for meat and hides, loss of habitat, and military strategies to deprive Native Americans of an essential food resource.29 Roosevelt knew the dwindling of the buffalo firsthand. He first visited the West in 1884, hunting buffalo in Dakota Territory, and returned re-



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peatedly. On a hunting trip on the Little Missouri River in 1887, he was disturbed to find buffalo hunted out, and he began to support their conservation (although he shot another one in 1899). A handful of zookeepers, ranchers, and private bison aficionados began to keep small herds of bison in captivity or in zoos. Led by the American Bison Society, bison were bred and released in protected areas, where they were managed. Surplus individuals were removed for the market or to establish other herds. Extinction was avoided, but it was a long haul, requiring a combination of captive breeding for eventual reintroduction and habitat conservation. Creation of Wind Cave National Park was part of this effort.30 But Wind Cave is much more than a national park: it is the site of the origin story of the Cheyenne Creek community on the Pine Ridge Indian Reservation of the Oglala Lakota tribe. As recounted to Sina Bear Eagle by Wilmer Mesteth, a tribal historian and spiritual leader, the story begins when animals and plants lived on the earth, but no bison and no people. People were living underground, waiting, as the earth was made ready for them. Through trickery by a spirit who lived on the surface of the earth, a few of these people, acting against the wishes of the Creator, were tempted aboveground. After a number of adventures, their unwillingness to work hard, and the arrival of a harsh winter, the Creator turned them into bison. Time passed and eventually the earth was ready for the remaining people, who passed through a passageway in the cave—Wind Cave—and arrived on earth. There they saw the hoofprints of a bison and were told by the Creator that everything they would need to survive would be provided by the bison.31 Bison were looked at longingly by cattle ranchers, who wanted to be able to breed cattle with the bison’s physiological capacities to survive on poor grass and endure harsh winters. They succeeded in a few cases, but in the process they introduced cattle genes into the genomes of some of the few surviving bison. From these animals the genes spread. The Wind Cave bison population appears to have escaped this fate, but many other bison herds across North America carry the genetic signature of cattle introgression. In 2005 the American Bison Society was reestablished. and one of its early meetings brought together stakeholders from Indigenous communities, the bison industry, NGOs, and landowners from throughout the

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range of the bison. This group developed a one-hundred-year vision for the ecological restoration of bison and identified a series of ways to increase their genetic diversity.32 In 2011, the American Bison Society convened a meeting of geneticists and bison managers, who discussed this genetic “contamination” of bison. They recognized that herds with no or little evidence of cattle-gene introgression should be a high priority for conservation, but they also recommended that management to eliminate cattle genes from bison should proceed with caution in order not to lose important bison genes. After all, many bison herds, with a low rate of cattle genes, continue to look like pure bison, and wallow, defecate, die, fight predators, and flee fires like bison. The group proposed that artificial (that is, human) selection should be avoided, even when aimed at purifying the genetic makeup of particular bison herds. In short, the group’s conclusion was both pragmatic and future orientated. It recognized the circumstances of bison conservation, and that their return from a historical population bottleneck had involved the introgression of some cattle genes. It accepted that a shading of cattle genes should not disqualify otherwise very bisonlike bison.33 As we have seen, technology is commonly thought of as the opposite of nature. When the grinding sound of a dirt bike interrupts a mountain hike, or when a siren cuts through the dawn chorus in a city park, both force into the present moment the disruptive quality of technology. But technology is much more complicated than is often thought. In ancient Greece it was thought that technology learned from or imitated nature. Today, “nature” is often available to people only through technologies— cross-country skis, watercraft, binoculars, GPS collars, automobiles, and so on. As we have written, nature is also often only still available thanks to the technologies of invasive-species management, visitor control, predator elimination, captive breeding, or fencing.34 The meanings of nature and technology shift according to time and place and are affected by experience, education, professional training, values, and political leanings. Historians Kranzberg and Pursell suggest that, “Technology, in a sense, is nothing more than the area of interaction between ourselves, as individuals, and our environment.”35 The complicated and elusive quality of synthetic biology is that it involves both nature and technology at the same time. Human intent shapes



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the tool, but the materials are, in most of contemporary synthetic biology, the building blocks of nature—base pairs, genes, and genomes. But in many ways, this is the story of conservation in general, the weaving of technology and nature—using one to save the other. Technology, like nature, is a social construct. As Williams wrote, “technology exists in reciprocity with both natural and social environments, all the while in symbiosis with its human creators. It has no logic of its own, and the politics that are inherent in every technology come from its human creators.” The politics of synthetic biology are as complicated as its technologies, and there is no agreement within communities of practice, within countries, or across the global community about how best to manage the astonishing powers that genetic editing confers.36 It is vain to wait for some sort of global consensus on the application of genetic tools. The genie is well out of the bottle in the biotech world. Genetically engineered organisms already exist not only in laboratories and factories, but in unbounded ecosystems, in eco, and more will join them. Some will be previously “wild” species, even if best known as pests and disease vectors. Others may be, perhaps, species whose genomes have been edited to increase their chances of survival. The decision to edit the genomes of wild species for conservation purposes is one that the conservation movement needs to face. The decision does not have to follow more general debates about genome editing in agriculture, biotechnology, or human health, although it will quite rightly be informed by them. The editing of wild species to save nature offers its own distinctive challenge. Its circumstances are different because of the urgency of the search for a response to human degradation of the biosphere. The nature that survives the twenty-first century is likely to be increasingly shaped by human action. Protected areas, fields, woodlands, urban forests, village greens, pipeline rights-of-way, and water impoundments will play dual roles, contributing to economy and society and yet providing space for nature. In all of them, even in remote tropical forest fastnesses or on distant mountains, there will be something novel about nature. Perhaps it will be species that have adapted to novel environments or that have been accidentally brought by people to live alongside those that evolved there. Or perhaps the frog calling from the forest pool will be genetically engineered, a mix of human choice and evolution’s blind

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selection by natural selection. Perhaps only frogs so modified by the tools of genome engineering will be left. In 1947, Aldo Leopold spoke at the newly created memorial to the passenger pigeon erected in the Wyalusing State Park in Wisconsin. The passenger pigeon was driven to extinction by a combination of hunting and habitat loss through the course of the nineteenth century. Leopold pointed out that it was a new thing for one species to mourn the extinction of another. He observed that the loss was not just of the species, but of its part in the wider ecosystem and in human culture, both the “seasonal changes on the Wisconsin landscape and the stories that went with that experience of the natural world.”37 Alan Holland and Kate Rawles wrote that conservation was about “negotiating the transition from past to future in such a way as to secure the transfer of maximum significance.” The diversity of the genome of each species, the diversity of the ecosystems of which they are part, are critical to the future of nature. In the language Aldo Leopold used, the first precaution of intelligent tinkering is to keep every cog and wheel. This is important not just to conservation but to human futures, to our understandings of the living beings with which we share the earth, and our understanding of ourselves.38 Conservation faces a huge challenge in the rise of genome editing and of biotechnology. There are threats and opportunities, risks and potential tools that could be used. There are already debates about genetic technologies in conservation, and these will surely grow. Indeed, they may grow more fierce and less tolerant. We could shy away from them and fall back on the things that are familiar. Or we could look forward to try to understand them and to respond to the challenges they raise. The prize is a future still shared by migrating cranes, wallowing bison, and wandering salmon. An earth whose diversity is living, flowing through and around us.

Appendix Scientific Names of Species

For the sake of readability, we have minimized the use of scientific names in the text of this book. However, not all the English names we use will be familiar, so here we give a list of the species we refer to, listed alphabetically by the common English name we use in the text, with their scientific names. This list is not exhaustive and is not intended to be in any sense definitive. We are naturalists not taxonomists. We recognize that scientific nomenclature is complex and evolving. We have used the scientific names with which we are familiar and apologize where these differ from those stipulated by appropriate authorities.

A African clawed frog Alewife Alfalfa Algerian mouse American bison American chestnut American cranberry American robin Amur leopard Anole lizard Apple, domestic Arabian oryx Ash, European Ash dieback fungus Asian elephant Asian long-horned beetle Asian rice

Xenopus laevis Alosa pseudoharengus Medicago sativa Mus spretus Bison bison Castanea dentata Vaccinium macrocarpon Turdus migratorius Panthera pardus orientalis Anolis spp. Malus domestica Oryx leucoryx Fraxinus excelsior Hymenoscyphus fraxineus Elephas maximus Anoplophora glabripennis Oryza sativa

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Asian tiger mosquito Atlantic horseshoe crab Atlantic killifish Atlantic salmon Auroch

Aedes albopictus or Stegomyia albopicta Limulus polyphemus Fundulus heteroclitus Salmo salar Bos primigenius

B Badger, European Bambara groundnut Banana (Cavendish) Barley Beaver, Eurasian Beaver, North American Bighorn sheep Bittern, Eurasian Black-faced honeycreeper (po‘ouli) Black-flanked rock wallaby Black-footed ferret Black rat Black redstart Black swallow-wort Blackcap Blue-and-yellow macaw Bluebell Bowerbird Breadfruit Bristlecone pine, Great Basin Brown-headed cowbird Brown rat Brown trout Brownbeard rice Brushtail possum Buriti palm Burrowing bettong Buzzard, common

Meles meles Vigna subterranea Musa acuminata, Cavendish subgroup Hordeum vulgare Castor fiber Castor canadensis Ovis canadensis Botaurus stellaris Melamprosops phaeosoma Petrogale lateralis Mustela nigripes Rattus rattus Phoenicurus ochruros Cynanchum louiseae Sylvia atricapilla Ara ararauna Hyacinthoides non-scripta family Ptilonorhynchidae Artocarpus altilis Pinus longaeva Molothrus ater Rattus norvegicus Salmo trutta Oryza rufipogon Trichosurus vulpecula Mauritia flexuosa Bettongia lesueur Buteo buteo

C California condor Cane toad Canola (rape) Carrot, domestic Catfish, channel

Gymnogyps californianus Rhinella marina Brassica napus Daucus carota sativus Ictalurus punctatus



Appendix: Scientific Names of Species 215

Cheatgrass (drooping brome) Bromus tectorum Cheetah Acinonyx jubatus Chestnut blight canker Cryphonectria parasitica Chicken Gallus gallus domesticus Chiffchaff Phylloscopus collybita Chinese chestnut Castanea mollissima Chinook salmon Oncorhynchus tshawytscha Chytrid fungus Batrachochytrium dendrobatidis and Batracho  chytrium salamandrivorans Codling moth Cydia pomonella Coho salmon Oncorhynchus kisutch Common heather Calluna vulgaris Corn (maize) Zea mays Cotton, upland or Mexican Gossypium hirsutum Cow Bos taurus Crab apple Malus sylvestris Crane, common or European Grus grus Creeping bentgrass Agrostis stolonifera Crown-of-thorns starfish Acanthaster planci

D Diamondback moth Dingo Domestic cat Domestic dog

Plutella xylostella Canis lupus dingo Felis catus Canis lupus familiaris

E Edible pea Eelgrass Elephant, African bush Elk Elm, English Ethiopian wolf Eucalyptus tree European ash European chestnut European honeybee European robin European viper (adder)

Pisum sativum Zostera spp. Loxodonta africana Cervus canadensis Ulmus procera Canis simensis Eucalyptus morrrisby Fraxinus excelsior Castanea sativa Apis mellifera Erithacus rubecula Vipera berus

F Fallow deer

Dama dama

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Appendix: Scientific Names of Species

Florida panther Fox, red Fruit fly, common

Puma concolor coryi Vulpes vulpes Drosophila melanogaster

G Gastric-brooding frog Rheobatrachus silus and Rheobatrachus   vitellinus Giant anteater Myrmecophaga tridactyla Giant armadillo Priodontes maximus Giant panda Ailuropoda melanoleuca Gorilla, mountain Gorilla beringei beringei Gray wolf Canis lupus Great white shark Carcharodon carcharias Gulf killifish Fundulus grandis

H Hobby, Eurasian Honey bee, European or Western Hops Horse, domestic House mouse Hyacinth bean

Falco subbuteo Apis mellifera Humulus lupulus Equus ferus caballus Mus musculus Lablab purpureus

I Iberian ibex Iberian lynx

Capra pyreanaica Lynx pardinus

J Jaguar Japanese or Korean chestnut Japanese knotweed Japanese macaque

Panthera onca Castanea crenata Fallopia japonica Macaca fuscata

K Kirtland’s warbler Koala Korean or Japanese chestnut

Setophaga kirtlandii Phascolarctos cinereus Castanea crenata

L Laughing owl Leopard

Sceloglaux albifacies Panthera pardus



Appendix: Scientific Names of Species 217

Lettuce Live oak Lynx, Eurasian

Lactuca sativa Quercus virginiana Lynx lynx

M Mammoth Maned wolf Marsh harrier Masai giraffe Millet, pearl Millet, finger Mink, European Monarch butterfly Morrisby’s gum Mosquito

Mammuthus spp. Chrysocyon brachyurus Circus aeruginosus Giraffa tippelskirchi Pennisetum glaucum Eleusine coracana Mustela lutreola Danaus plexippus Eucalyptus morrisbyi Anopheles spp. and Aedes spp.

N New World screwworm fly Nightingale Northern giraffe Northern quoll

Cochliomyia hominivorax Luscinia megarhynchos Giraffa camelopardalis Dasyurus hallucatus

O Oak tree, English Ocean pout ¯ hi‘a lehua tree ‘O Okapi Orange, common Orangefoot pimpleback mussel Orangutan Oxlip

Quercus petraea Zoarces americanus Metrosideros polymorpha Okapia johnstoni Citrus sinensis Plethobasus cooperianus Pongo spp. Primula elatior

P Palmer amaranth Panamanian golden frog Panda, giant Papaya Passenger pigeon Pig, domestic Pink bollworm Pink fairy armadillo Pleistocene cave bear

Amaranthus palmeri Atelopus zeteki Ailuropoda melanoleuca Carica papaya Ectopistes migratorius Sus scrofa domesticus Pectinophora gossypiella Chlamyphorus truncatus Ursus spelaeus

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Polynesian rat Potato Prairie dog Proboscis monkey Puma Purple hairstreak butterfly Pyrenean ibex

Rattus exulans Solanum tuberosum Cynomys spp. Nasalis larvatus Puma concolor Neozephyrus quercus Capra pyrenaica pyrenaica

Q Quagga Quaking aspen

Equus quagga Populus tremuloides

R Raccoon dog Rainbow trout Red fox, European Red peppers Rhea, greater Rice, Asian Rodrigues fody Rosebay willow herb Rough fatmucket mussel

Nyctereutes procyonoides Oncorhynchus mykiss Vulpes vulpes Capsicum annuum Rhea americana Oryza sativa Foudia flavicans Chamaeneerion angustifolium Lampsilis straminea

S Sabre-toothed cat Sad elliptio mussel Sea beet Sea lamprey Sockeye salmon Sooty tern Sorghum South Georgia pipit Southern giraffe Southern house mosquito Soybean or soya bean Sparrow, house Sparrow, Italian Sparrow, Spanish Sparrow, Turkestan house Spotted knapweed Stoat Streamer-tailed tyrant

Smilodon fatalis Elliptio lugubris Beta vulgaris ssp. maritime Petromyzon marinus Oncorhynchus nerka Onychoprion fuscatus Sorghum bicolor Anthus antarcticus Giraffa giraffa Culex quinquefasciatus Glycine max Passer domesticus Passer italiae Passer hispaniolensis Passer domesticus bactrianus Centaurea stoebe Mustela erminea Gubernetes yetapa



Appendix: Scientific Names of Species 219

Sugar beet Sugarcane Sweet potato Sweet wormwood

Beta vulgaris ssp. vulgaris Saccharum officinarum Ipomoea batatas Artemisia annua

T Tasmanian devil Teff Thylacine or Tasmanian tiger Tilapia, Nile Tobacco Tristan albatross Tsetse fly Tuatara

Sarcophilus harrisii Eragrostis tef Thylacinus cynocephalus Oreochromis niloticus Nicotiana tabacum Diomedea dabbenena Glossina spp. Sphenodon punctatus

U ——

V Varroa mite

Varroa destructor

W Water hemp Amaranthus rudis and Amaranthus   tuberculatus Western corn rootworm Diabrotica virgifera virgifera Wheat, bread Triticum aestivum White button mushroom   or champignon Agaricus bisporus White-footed mouse Peromyscus leucopus White-tailed deer Odocoileus virginianus Whooping crane Grus americana Wild garlic Allium ursinum Wildcat, European Felis silvestris Wildebeest, common or blue Connochaetes taurinus Wolf, Gray Canis lupus Wolf, Mexican Canis lupus baileyi Wolf, Mongolian Canis lupus chanco Wood anemone Anemone nemorosa Woolly mammoth Mammuthus primigenius

X ——

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Appendix: Scientific Names of Species

Y Yak, domesticated Yellow-crested cockatoo Yellow fever mosquito Yew, English

Bos grunniens Cacatua sulphure Aedes aegypti Taxus baccata

Z Zebra, plains Zebra fish Zebra mussel

Equus quagga Dani rerio Dreissena polymorpha

Notes

1. The Place of Nature 1. Ray, J. (1660), Catalogus plantarum circa Cantabrigiam nascentium [Catalogue of Cambridge plants], ed. Ewen, A. H., and Prime, C. T., 1975, Wheldon and Wesley, Hitchin, UK; Hopkins, G. M. (1918), “The May Magnificat,” in Gerard Manley Hopkins: The Major Works, ed. C. Phillips, Oxford University Press, Oxford. 2. Rackham, O. (1975), Hayley Wood: Its History and Ecology, Cambridge and Isle of Ely Naturalists’ Trust, Cambridge. 3. Rackham, O. (1976), Trees and Woodlands in the British Landscape, J. M. Dent, London; Rackham, O. (1986), The History of the Countryside, J. M. Dent, London. 4. Lankester, E. R. (1914), Nature reserves, Nature, March 12: 33–35 (p. 33). 5. On British conservation, see Sheail, J. (1976), Nature in Trust: The History of Nature Conservation in Great Britain, Blackie, Glasgow. 6. Protected Planet, World Database of Protected Areas, https://www.protected planet.net, accessed June 2019; Aerts, R., Van Overtveld, K., November, E., et al. (2016), Conservation of the Ethiopian church forests: Threats, opportunities and implications for their management, Science of the Total Environment 551–552: 404–414; Rezende, C. L., Scarano, F. R., Assad, E. D., et al. (2018), From hotspot to hopespot: An opportunity for the Brazilian Atlantic Forest, Perspectives in Ecology and Conservation 16: 204–214. 7. Stevenson Center (n.d.), Adlai E. Stevenson II, https://stevensoncenter.org /about/stevenson.php, accessed April 8, 2020; Buckminster Fuller, R. (1968), Operating Manual for Spaceship Earth, Southern Illinois University Press, Carbondale (2008 ed. Lars Muller Publishers, Zurich); Höhler, S. (2015), Spaceship Earth in the Environmental Age 1960–1990, Taylor and Francis, London. 8. Adams, W. M. (2020), Green Development: Environment and Sustainability in a Developing World, Routledge, London.

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NOTES TO PAGES 200–208 263

17. Sandler, R. (2011), Is artefactualness a value-relevant property of living things?, Synthese 185: 89–102. 18. Sandler, R., The ethics of genetic engineering; Palmer, C. (2016), Saving species but losing wildness: Should we genetically adapt wild animal species to help them respond to climate change?, Midwest Studies in Philosophy 40: 234–251. 19. Sarkar, S. (1999), Wilderness preservation and biodiversity conservation— Keeping divergent goals distinct, BioScience 49: 405–412. 20. Hobbs, R. J., Higgs, E. S., Hall, C. M., eds. (2013), Novel Ecosystems: Intervening in the New World Order, Wiley-Blackwell, Chichester, UK. 21. See, for example, Smith, M. (2019), (A)wake for “the passions of this earth”: Extinction and the absurd “ethics” of novel ecosystems, Cultural Studies Review 25: 119–134; Marris, E. (2011), Rambunctious Garden: Saving Nature in a Post-Wild World, Bloomsbury Press, New York; McEuen, A. B., and Styles, M. A. (2019), Is gardening a useful metaphor for conservation and restoration? History and controversy, Restoration Ecology 6: 1194–1198. 22. Smith, B., Bradshaw, C. J. A., Richie, E., et al. (2019), The dingo is a true-blue, native Australian species, The Conversation March 6, http://theconversation.com/, accessed August 2019; Townsend, A. K., Staab, H. A., and Barker, C. M. (2019), Urbanization and elevated cholesterol in American crows, The Condor 121: 1–10. 23. This problem is carefully discussed by Shapiro, B. (2015), How to Clone a Mammoth: The Science of De-Extinction, Princeton University Press, Princeton, NJ; Seddon, P. J. (2017), The ecology of de-extinction, Functional Ecology 31: 992–995. 24. Lorimer, J., Sandom, C., Jepson, P., et al. (2015), Rewilding: Science, practice, and politics, Annual Review of Environment and Resources 40: 39–62; Perino, A., Pereira, H. M., Navarro, L. M., et al. (2019), Rewilding complex ecosystems, Science 364 (6438): eaav5570 (2019); Taylor, P. (2005), Beyond Conservation: A Wildland Strategy, Earthscan, London. 25. Child, M. F., Selier, S. A. J., Radloff, F. G. T., et al. (2019), A framework to measure the wildness of managed large vertebrate populations, Conservation Biology 33: 1106–1119. 26. Lorimer, J. (2017), Probiotic environmentalities: Rewilding with wolves and worms, Theory, Culture & Society 34: 27–48. 27. Elliot (1982), Faking nature, 87. 28. Bell, D. A., Robinson, Z. L., Funk, W. C., et al. (2019), The exciting potential and remaining uncertainties of genetic rescue, Trends in Ecology & Evolution 12: 1070–1079; Funk, W. C., Forester, B. R., Converse, S. J., et al. (2018), Improving conservation policy with genomics: A guide to integrating adaptive potential into U.S. Endangered Species Act decisions for conservation practitioners and geneticists, Conservation Genetics 20: 115–134; Smith, T. B., Kinnison, M. T., Strauss, S. Y., et al. (2014), Prescriptive evolution to conserve and manage biodiversity, Annual Review of Ecology, Evolution, and Systematics 45: 1–22; Eizaguirre, C., and Baltazar-Soares, M. (2014), Evolutionary conservation—Evaluating the adaptive potential of species, Evolutionary Applications 7: 963–967.

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29. Isenberg, A. C. (2001), The Destruction of the Bison: An Environmental History, Cambridge University Press, Cambridge. 30. Jeffers, H. P. (2003), Roosevelt the Explorer: Teddy Roosevelt’s Amazing Adventures as a Naturalist, Conservationist, and Explorer, Taylor Trade Publishing, Lanham, NY. 31. The Lakota Emergence Story, US National Parks Service, available at: https:// www.nps.gov, accessed September 2019. 32. Sanderson, E. W., Redford, K. H., Weber, B., et al. (2008), The ecological future of the North American bison: Conceiving long-term, large-scale conservation of wildlife, Conservation Biology 22: 252–266; Redford, K. H., Amato, G., Baillie, J., et al. (2011), What does it mean to successfully conserve a (vertebrate) species?, BioScience 61: 39–48. 33. Recommendations from the American Bison Society Meeting on Bison Ecological Restoration (2011), Tulsa, Oklahoma, March 23–25, unpublished. 34. Frannsen, M., Lokhorst, G.-J., Van de Poel, I. (2018), Philosophy of technology, in The Stanford Encyclopedia of Philosophy, ed. Zalta, E. N., available at: https:// plato.stanford.edu/, accessed April 2020. 35. Melvin Kranzberg and Carroll Pursell (1967), quoted in Williams, J. C. (2010), Understanding the place of humans in nature, in The Illusory Boundary: Environment and Technology in History, ed. Reuss, M., and Cutcliffe, S. H., University of Virginia Press, Charlottesville, VA, 9–25 (p. 16). 36. Williams, Understanding the place of humans in nature, 21. 37. Enright, K. (2019), Exhibiting extinction: Martha and the monument, two modes of remembering nature, Cultural Studies Review 25: 154–171; Jørgensen, D. (2019), Recovering Lost Species in the Modern Age: Histories of Longing and Belonging, MIT Press, Cambridge, MA. 38. Holland, A., and Rawles, K. (1993), Values in conservation, ECOS: A Review of Conservation 14 (1): 14–19; Leopold, A. (1953), Round River: From the Journals of Aldo Leopold, ed. Leopold, L. B., Oxford University Press, New York.

Index

Adams, Bill, x–xi, xii, 1, 4, 32, 41 Africa: agricultural use of synthetic biology in, 119–20; Congo Basin, 14, 39; Gough Island, 141; Maasai Mara, 4; mosquito and malaria control in, 131; Okavango Delta, 39; regulation of synthetic biology in, 179; Selous, 4; Serengeti, 4 African clawed frog, 93 African Orphan Crops Consortium, 119 Agricultural (Neolithic) Revolution, 2, 3, 35–36, 53, 78 agriculture: alteration of genomes of domestic organisms, 17; ancient woodlands/industrial agricultural landscape, juxtaposition of, 1–4, 9, 13; antibiotic use and resistance, 80; in cerrado biome, Brazil, 46–47; cisgenesis and transgenesis, use of, 95; elephant conservation management and, 32–33; “escape” of novel genetic devices into nature, problem of, 125; fertilizers, artificial, 12, 120; gene editing technologies, advent of, 119; harvesting, genetic effects of, 77–78; organic farming and gene flow, 126; selective breeding practices and, 91–93. See also domestic crops and animals, genetically modified (GM) crops and organisms

Agrobacterium tumefaciens, 94, 99 alewife, 204 algae, x, 15, 59, 110, 121, 123–24, 144, 153, 155, 156, 159 Alien Resurrection (film), 165 alleles, 62 Alliance for a Green Revolution in Africa, 119 Amazon/Amazonia, 14, 46, 47, 116 Amend, Anthony, 76 American Bird Conservancy, 147 American bison, 14, 15, 82, 208–10 American Bison Society, 208–10 American chestnut, 136–39, 149, 165–66, 204 American Chestnut Foundation, 138 American Samoa, coral reefs of, 155 Amino Labs, 113 amphibians, disease-related global decline of, 149–52 Amur leopard, 14–15 Amyris, 110 ancient woodlands, 1–4, 9 Anders, Bill, 5 animal welfare ethics, 165 animal-machine hybrids, 104–5, 113–14 Annihilation (VanderMeer), 168 Antarctica, 7, 13, 25, 32, 36, 49 anthromes versus biomes, 6 Anthropocene, 8, 40, 193

265

266

INDEX

Anthropogenic climate change, 36, 37 antibiotics, microbial resistance to, 80 anti-CRISPRs, 101 ants, 145 Apollo 7, 5 apples/apple trees, 127 aquaculture. See fish and fisheries AquAdvantage salmon, 96, 108, 129 Arabian oryx, 82 Arabidopsis thalania, 66 ArborGen, 139 Arctic, 39, 41, 81, 84, 186 Argentina: GM crops in, 47, 109; herbicide resistance in, 127; pink fairy armadillo of, 49; regulation of synthetic biology in, 179; Aristotle, 70 Arkansas National Wildlife Refuge, 21 artemisinin, 110–11 artificial light, evolutionary responses to, 79 the artificial versus the natural, xi, 20–23, 42–44, 194–97, 207 Ascension Island, invasive/feral species on, 148 asexual reproduction, 61 ash dieback disease, 85, 149 Asian elephants, 163, 205 assisted evolution, 154–55 Atlantic killifish, 85 Atomic Energy Commission, US, 92 Atwood, Margaret, Oryx and Crake, 167–69 aurochs, 53, 160–61 Australasian pteropid fruit bats, 145 Australia: colonization of, 35, 143; “command and control” approach to conservation management in, 30, 32; DIYbiosphere Project in, 113; dsRNA used in, 123; emergence of life, evidence for, 49; genetic conservation in, 83, 85, 87; glyphosate resistance in, 127; GM canola in, 126, 156; Great Barrier Reef, 76, 85, 153; invasive/ feral species in, 37, 143, 144, 226n30; organic farming and gene flow in, 126;

public health controls, use of synthetic biology in, 183; regulation of synthetic biology in, 179. See also specific animals unique to Australia Australian jumping spider, 52 automation of synthetic biology technologies, 102–3 Aventis, 10 avian malaria and avian pox virus, 146   Bacigalupi, Paolo, The Windup Girl, 169 back-breeding, 160–61 bacteria, 15–16, 47–49, 61, 67, 74–76, 80, 89, 93–94, 99, 101–6, 110, 14, 117, 121–23, 128, 150–53, 156–59 Ballard Locks Fish Ladder, Seattle, WA, 191 bananas and banana leaf wilt, 115, 122 Bangladesh: Golden Rice in, 120, 243n37; regulation of synthetic biology in, 179 barley, 2, 57 Bateson, William, 56 bats, 145, 149 Bayer, 139 Beagle (ship), 53–54 beavers, 41, 67 beet crops, 127 bighorn sheep, 68–70, 72, 73, 82 Bill and Melinda Gates Foundation, 119 BioBricks, 103, 113 BioBuilder Educational Foundation, 114 biocrusts, 157 biodiversity, 13–17, 45–67; in ancient woodlands, 2; of cerrado biome, 46–48; commercial/practical applications of synthetic biology, as resource for, 114–17; defined, 16; DNA, dependent on dynamism of, 67; of ecosystems, 13–14, 16; evolution of, 48–55, 61–65; genes and, 55–68 (See also genes); genetic intervention and, 191–94; genomes and, 16–17, 65–67 (See also genome); historical study of, 50–56; hybridization diminishing, 73; importance to proper functioning of ecosystems, 48; loss of, ix, xi, 8, 39, 145;



of microbiome, 15–16, 47, 49–50; of species, 14–15, 16, 47, 50 bioeconomy, 111–14 biofuels, 109–10, 158 biolistic particle delivery system (“gene gun”), 94, 96 biomes versus anthromes, 6 bioprospecting, 116 bison. See American bison bittern, 194 black sigatoka virus, 115 black swallow-wort, 38 blackcap warbler, 1 black-flanked rock wallaby, 83 black-footed ferret, 34, 82, 140, 198 Blue Planet image, 5 Blue Ridge, Virginia, 136 blue-and-yellow macaw, 47, 64 bluebells, 1 Bock, Ralph, 110 Bolivia, regulation of synthetic biology in, 179 bollworm, 122 Bolt Threads, 109 Bonn Challenge, 34 Borlaug, Norman, 119 Borneo rain forests, 14 BovB, 76 bowerbirds, 49 Boyer, Herbert, 93 Branson, Richard, 108 Brazil: Amazon/Amazonia, 14, 46, 47, 116; Atlantic Forest, 4; cerrado biome, 46–48, 50, 51, 61, 64, 102; de novo domestication of wild tomato in, 115; Emas National Park, 46–48; giant anteaters in, xi, 45–47; glyphosate resistance in, 127; GM crops in, 109; Oxitec mosquito releases, 130; Pantanal, 13, 39; reforestation commitments in, 34; regulation of synthetic biology in, 179 Bronx Zoo, NYC, 137 Brookhaven National Laboratory, 137 brushtail possum, 32, 143 Bt cotton and maize, 109, 122, 128

INDEX 267

buffalo. See American bison buriti palm, 47 burrowing bettong 83 buzzard, 204   C3 BIOTECH, 110 Calgene, 12 California condor, 60, 67, 82 Cambridgeshire (UK), ancient woodlands/ industrial farmland of, 1–4 Canada: AquAdvantage salmon sold in, 108; beaver, sequencing genome of, 67; DIYbiosphere Project in, 113; GM crops in, 109; Kirtland’s warblers in, 31; regulation of synthetic biology in, 179; white nose syndrome in bats of, 149; whooping cranes in, 20; Wood Buffalo National Park, 20 cane toads, 87, 143 Canidae. See specific types canola, 2, 11, 18, 109, 126, 156, 258n13 Cargill, 108 Carlson, Rob, 14, 104, 111, 126, 127 carp, 108 Carson, Rachel, 14, 104, 111, 126–27; Silent Spring, 12, 43, 196 Cartagena Protocol on Biosafety, 176, 177 cassava, 120 catfish, 108 cats, domestic: cloning, 161; hybridization with European wildcats, 84; as invasive/ feral species, 32, 143, 148 cattle/cows: biodiversity and, 53; commercial/practical applications of synthetic biology and, 107, 108; genetic conservation and, 76; Heck cows, 160–61, 162; Highland cattle, 41, 42; methane production by, 158–59; nature and conservation issues, 8, 15–16, 41, 42, 47, 209, 210; synthetic biology and, 91, 93 cave bears, 83 Cavendish banana, 115 CBD (Convention on Biological Diversity), 16, 35, 72, 116, 140, 176–77 Celera Genomics, 66

268

INDEX

“cell-free” synthetic biology, 103–4 cerrado biome, Brazil, 46–48, 50, 51, 61, 64, 102 cesium 137, 186 Chakrabarty, Ananda, 116 cheatgrass, 153 chemical mutagens, 92–93 Chernobyl nuclear disaster and site, 39, 63, 186–87 chestnut blight canker, 137, 138 chestnut trees, 136–39, 149, 165–66, 204 chickens, 111, 161 chiffchaff warbler, 1 Chin, Jason, 100–101 China: Bt cotton, problems with, 128; commercial/practical applications of synthetic biology in, 47, 90, 111–14, 118, 122, 126, 128, 175; CRISPR-baby scandal in, 118; gene flow between domestic and wild rice in, 126; reforestation efforts in, 34; regulation of synthetic biology in, 179 Chinese Academy of Chinese Medical Sciences, 110 Chinese Academy of Sciences, 120 Chinese chestnut, 136, 137–38 chipmunks, 85 church forests, Ethiopia, 4 chytridiomycosis and chytrid fungus, 149–52 cisgenesis, 95 CITES (Convention on International Trade in Endangered Species), 72, 176 citrus greening, 122 classification of species, 68–74 climate change: Anthropogenic, 36, 37; coral reef conservation and, 153–55; deliberate adaptive introgression and, 85; disease spread and, 146; ecological response to, 35–37; ecological restoration and, 157–59; synthetic biology and, 90 Clinton, Bill, 66 cloning, 161–62 clothing, 109 codling moth, 93 Cohen, Stanley, 93

colchicine, 92 Cold War and the Space Race, 5 commercial/practical applications of synthetic biology, 107–35; bioeconomy of, 111–14; common products, development of, 107–11; conception of genes, economic associations of, 117; in domesticated crops and animals, 118–21; “escape” into nature, problem of, 117–18, 123–25 (See also in eco synthetic biology); nature/biodiversity as resource for, 114–17; novel life-forms, 133–34; pesticides, herbicides, and insecticides, 121–23; popular/student involvement in, 113–14; terraforming, 134; transgenic American chestnut and, 139 community genetics, 64–65 Congo Basin, Africa, 14, 39 conservation genetics, 16 Conservation International, 48 conservation management, 20–44; artificial conservation techniques, use of, xi, 20–23, 42–44; climate change and, 35–37; “command and control” approach to, 29–34; concept of nature and, 23–27, 42–44; equilibrium paradigm, breaking, 27–29; evolutionary conservation, 67; historical development of, 24–28; in human-created environments, 37–40; of invasive/feral species, 32, 37–38; protected areas model for, 1–4; remaining wild lands, priority given to, 7–8; restoration of ecology as goal of, 34–35, 36, 41, 201–2; rewilding movement, 40–42. See also conservation using synthetic biology; genetic conservation conservation using synthetic biology, ix–xii, 17–19, 136–66; American chestnut restoration plan, 136–39, 149, 165–66; biodiversity issues, 191–94; climate change mitigation, 153–55, 157–59; concerns regarding, 139–40, 147–49, 151–52, 163–65, 172–73, 187–88, 196–201; coral reefs and climate change, 153–55; CRISPR, 142, 143, 144, 149, 150, 155; de-extinction



(recreation of extinct species), 159–66, 169, 205; disease control, 140, 145–47, 149–53; in eco synthetic biology, concerns of conservationists regarding, 134–35; ecological restoration, 157–59; engineered gene drives, 142–49 (See also engineered gene drives); fungal diseases, 149–52; gene editing tech­ nologies, 138, 144, 152–53; growing strangeness of nature and, ix, x, 205; invasive/feral species, control or elimination of, 141–45, 153; microbiome manipulation, 152–53; product substitution, 155–57; RNA silencing/ RNA interference (RNAi), 145. See also naturalness, synthetic biology, and conservation conservation-reliant species, 31 Convention on Biological Diversity (CBD), 16, 35, 72, 116, 140, 176–77 Convention on International Trade in Endangered Species (CITES), 72, 176 Convention on the Law of the Seas., 176 corals and coral reefs, 76, 85, 144, 153–55, 188 corn/maize: biodiversity issues, 53, 62; commercial/practical applications of synthetic biology and, 109, 119, 120–23, 126, 128; conservation using synthetic biology and, 138; nature and conservation issues, 9–11, 13; synthetic biology and, 91, 92, 95 Correns, Carl, 56 Corteva Agriscience, 122 cosmetics and scents, 110 cotton, 11, 109, 122, 126, 128 Court of Justice of the European Union, 179 COVID-19 pandemic, 145, 170 cowbirds, 31 cows. See cattle/cows coyotes, 202 crab apple trees, 127 cranes, 20–22, 194–95 creeping bentgrass,127 Crick, Francis, 56 Criminal Damage Act of 1971 (UK), 10

INDEX 269

CRISPR: commercial/practical uses of, 104, 108, 111, 113, 114, 115, 118, 122, 123; for conservation purposes, 142, 143, 144, 149, 150, 155; debates about use of, 198, 199; development of, 99–101; gene drives, creating, 129, 131, 132–33; government regulation of organisms edited by, 178; human embryos, used on, 118; transformative power of, ix–x CRISPR/Cas9 and CRISPR/Cpf1, 142 crown-of-thorns starfish, 144 crows, 202 cryptorchidism, 69 cultured foods, 107–9, 255n62 cyanobacteria, 157, 159 Czechoslovakian wolfdog, 160   dairy products, 108 “daisy-chain” gene drives, 132, 181 DARPA (Defense Advanced Research Projects Agency), US, 123 Darwin, Charles, 52–55, 61, 77; “On the Tendency of Species to Form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection” (with Wallace), 54; On the Origin of Species, 54 data-storage unit, using DNA as, 104 de Vries, Hugo, 56 debates about synthetic biology, 167–90; conservation using synthetic biology, 139–40, 147–49, 151–52, 163–65, 172–73, 187–88, 196–201; CRISPR, 198, 199; de-extinction, 163–65, 169; degree of care, questions about, 187–90; different applications, treatment of, 180–82; in eco synthetic biology, 117–18, 123–25, 131–32, 134–35; engineered gene drives, 147–49, 171, 181; gene editing, 170, 197; genome editing, 176, 181, 182, 185, 188, 189; GM crops and organisms, 9–13, 120, 174; in literature, 167–69; possibilities matched by concerns, 169–70; power, authority, and control issues, 168–69, 174–77; precautionary principle/ approach, 182–83; public trust and

270

INDEX

debates about synthetic biology (cont.) engagement, 183–87; scientific and public concerns, 170–74; “Trojan horse” fears, 139, 171, 257n8. See also regulation of synthetic biology deer, white-tailed 203 de-extinction (recreation of extinct species), 159–66, 169, 205 Defense Advanced Research Projects Agency (DARPA), US, 123 deliberate adaptive introgression, 85–86 dengue fever, 130, 183 Denisovans, 74 design-build-test-learn cycle, 101–2 devil facial tumor disease, 145 diamondback moth, 17–18 dingo, 202 dinosaurs, 49, 160 disease: antibiotics, microbial resistance to, 80; climate change and, 146; conservation use of synthetic biology to control, 140, 145–47, 149–53; deliberate adaptive introgression as means of combating, 85; fungal diseases, 149–52; mosquito vectors, 130–33, 146–47; protection of wild animals against, 34; zoonotic, 145–46. See also specific diseases disruptive technologies, 112 diversity, genetic. See biodiversity DIYbiosphere Project, 113 DNA (deoxyribonucleic acid): additional non-biological bases, inserting, 105, 134; “barcodes,” species identification via, 86; biodiversity dependent on dynamism of, 67; changes to, 62–64; as data-storage unit, 104; discovery and structural analysis of, 55–57; environmental DNA/eDNA, 86; “Frozen Zoo,” San Diego Zoo, 81; functions of, 57–59; gene drives, 129; hachimoji DNA and RNA, 105; horizontal gene transfer of, 75; human hybridization revealed by, 74; “junk” DNA, 75, 193; location[s] of, 57; radiation and chemicals, deliberately introducing mutations through, 92–93, 137; recombi-

nant DNA technologies, 93–96; RNA and, 59; sequencing, 65, 66, 177; traits expressed by, 59–60, 61. See also gene editing; synthetic biology DNA nanotechnology, 104 DNA origami, 104 dogs, domestic: back-breeding, 160; cloning, 161; feral, 39; poison bait as threat to, 32, 141–42; rabies and, 34; selective breeding of, 90–91; wolves, cross-breeding with, 84 Dolly the sheep, 161, 165 domesticated crops and animals: agri­ cultural alteration of genomes of, 17; assisted evolution compared to domestication, 154; commercial/ practical applications of synthetic biology in, 118–21; creation of, 53; engineering pest resistance in, 47, 121–22, 126–27; gene flow/genetic transfer or introgression, 125–29; methane pollution and, 158–59; microbes, domestication of, 91; rewilding movement and traditional domestic breeds, 41, 42; selective breeding of, 90–93, 95, 115. See also cats; dogs; pesticides, herbicides, and insecticides double-stranded RNA (dsRNA), 123 Doudna, Jennifer, 100 dsRNA (double-stranded RNA), 123 DuPont, 94 Dutch elm disease, 61, 149   E. coli (Escherichia coli), 63, 93 Earth BioGenome Project, 67, 115 Ebola, 145 ecological restoration, 157–59 economics of synthetic biology. See commercial/practical applications of synthetic biology EcoRI, 93 ecosystem: concept of, 13–14, 16, 27–28; de-extinction critique and, 163–64 ecosystem microbiomes, 76 eDNA (environmental DNA), 86 elephant ivory, synthetic, 156–57



elephants, 32–33, 163, 205 elk, 15 Ellen MacArthur Foundation, 8 Elliott, Robert, 200 elm trees, 61, 149 Elwha and Glines Canyon Dams, WA, removal of, 192 Emas National Park, Brazil, 46–48 Emerging Risks Team, Lloyds of London, 90 Endangered Species Act (US), 31, 72, 165 endemic species, 47, 48 Endy, Drew, 103 engineered gene drives, 129–33; concerns about, 147–49, 171, 181; conservation dangers of, 147–49; CRISPR used to create, 129, 131, 132–33; “daisy-chain”/ “localized”/“locally fixed allele,” 132, 148, 181; disease controls in endangered species, 145–47; invasive species, controlling/eliminating, 142–45; pesticides, herbicides, and insecticides, 130–31, 148; regulatory issues, 132, 181; reversal gene drives, 132, 148, 181 engineering heritage of synthetic biology, 97–98 Enlightenment, 23–24 enteroviruses, 111 environmental DNA (eDNA), 86 environmental influences on genes, 59, 64 Environmental Protection Agency, US, 128 epigenetics, 63–64 equilibrium paradigm in ecology, 27–29 “escape” into nature, problem of, 117–18, 123–25. See also in eco synthetic biology Escherichia coli (E. coli), 63, 93 ETC Group (Action Group on Erosion, Technology and Concentration), 135, 181 Ethiopian wolf, 34 eukaryotes, 57–59, 61, 66, 67, 115 European chestnut, 136 European Commission, 182 European Food Safety Agency, 178, 179 European Union (EU): DIYbiosphere Project in, 113; GM crops and organ-

INDEX 271

isms, 11, 127, 174, 178–79; regulation of synthetic biology in, 178–79 European viper, 83 European wildcat, 84 evolution, 48–55, 61–65, 77–80, 206–8 evolutionary conservation, 67 evolutionary rescue, 82–83 ex situ conservation, 80–82 extinction: de-extinction (recreation of extinct species), 159–66, 169, 205; as evolutionary process, 49, 50; through hybridization/introgression, 83–84   fabric, 109 farming. See agriculture feral species, 32, 39, 42, 143–48 fertilizers, artificial, 12, 120, 128 Finless Foods, 108 Firebird Biomolecular Sciences, 105 fish and fisheries: gene flow/genetic transfer or introgression, 128–29; genetic effects of commercial fishing, 77–78; genetic rescues, 83; hybridization due to human activities, 85; invasive fish, 144–45; marine predators, 8; product substitution, 155–56; synthetic fish, as food, 108. See also marine environments; specific types of fish Fish and Wildlife Service, US, 21–22, 31, 34, 60, 140, 147 fish oil market, 155–56 flamingos, 69 Flavr Savr tomato, 12, 108–9 Flores, Fernando, Tears of the Trufflepig, 169 Florida panthers, 68–70, 72, 73, 82, 83, 87 Food and Drug Administration, US, 178 foodstuffs: GM crops and organisms, 9–13, 17, 23, 47, 95–96, 108–9; synthetic/cultured, 107–9. See also specific types forage fish, 155–56 Forest Service, US, 31 forests. See trees and forests “fourth industrial revolution” (genetic engineering), 112, 114

272

INDEX

Frankenstein (Shelley), 12–13, 159, 160, 167–69, 196 Frankenstein foods/Frankenfoods, 12–13 Franklin, Rosalind, 56 freshwater environments, 52, 141, 144–45, 191 freshwater mussels, 51–52 “Friendly” (Oxitec mosquitoes), 130 Friends of the Earth, 135 frogs, 82, 93, 149–52, 160, 207, 211–12 “Frozen Zoo,” San Diego Zoo, 81, 162 fruit flies, 66, 73, 93, 130 Fukushima Daiichi nuclear disaster, Japan, 39 fungal diseases, 149–52 future naturalness, 39–40   Galápagos Islands, 53, 61 Garza, John Carlos, 83 gastric-brooding frog, 160 Gates, Bill, 108, 119, 181 GBIRd (Genetic Biocontrol of Invasive Rodents), 142 gene drives, engineered. See engineered gene drives gene editing technologies, 96–101; in agriculture, 119; aquaculture, use in, 108; conservation uses of, 138, 144, 152–53; de-extinction and, 160, 162–63; naturalness in conservation and, 208; regulation of, 179, 181–82; risks of/ debates about, 170, 197; RNA used in, 104; standardized steps for, 98–99; transformative power of, ix–x, 17, 100–101; wide application of, 19. See also specific technologies gene expression, 63–64 gene flow/genetic transfer or introgression, 125–29, 209–10 “gene gun” (biolistic particle delivery system), 94, 96 General Electric, 116 genes, 55–68; concept of, 59–60; defined, 58–59; ecosystem/landscape level, change at, 64–65; environmental influences on, 59, 64; epigenetics, methylation, and gene expression, 59,

63–64; expression of, 60, 97; genetic conservation and analysis of, 86–87; mutation of, 62–63; origins of concept of, 55–56; RNA (ribonucleic acid), 57–59, 63; species, crossing between, 74–77. See also DNA Genetic Biocontrol of Invasive Rodents (GBIRd), 142 genetic bottlenecks, 60, 68–69 genetic conservation, 68–88; analysis of genes and genomes, 86–87; classification of species and, 68–74; deliberate adaptive introgression, 85–86; DNA “barcodes,” species identification via, 86; environmental DNA/eDNA, 86; ex situ conservation, 80–82; genomes, transfer of genes between, 74–77; human environments, effects of, 77–80; hybridization and, 70, 73–74, 77, 83–85; inbreeding and low genetic diversity, managing, 68–69, 73, 82–83 genetic rescue, 82–83 genetic variation and speciation, 61–65 genetically modified (GM) crops and organisms, 17, 23, 47; commercial uses of synthetic biology and, 108–9, 111, 120; conservation uses of synthetic biology and, 138–39; controversies over, 9–13, 120, 174; development of synthetic biology and, 95–96; gene flow/genetic transfer or introgression and, 125–29; regulation of, 178–79, 181 Genome 10K project, San Diego Zoo, 67 genome editing, 98–101; commercial/ practical applications, 108, 115, 123, 129; in conservation, 138–41, 144, 151, 157; debates about, 176, 181, 182, 185, 188, 189; naturalness issues and, 193, 195–96, 198, 200, 205, 211, 212 genomes, 65–67; biodiversity of, 16–17; defined, 65; genetic conservation and analysis of, 86–87; sequencing of, 65–67; transfer of genes between, 74–77 genomics, 119 genotoxic chemicals, 63



Germany: Berlin, urban ecosystems in, 37; Bonn Challenge, 34; Clean Air Act (1974), 182 giant anteaters, xi, 45–47 giant armadillo, 47 giant ground sloths, 50 “giant terror birds,” 50 Gingko Bioworks, 102, 110 Ginsberg, Alexandra Daisy, 133 giraffes, 71 Global Justice Ecology Project, 139 GloFish, 96 glyphosate and glyphosate resistance, 47, 126–27 glyptodonts, 50 GM crops and organisms/GMOs. See genetically modified (GM) crops and organisms goats, genetically engineered, 109, 111 Golden Rice, 120 Gore, Al, An Inconvenient Truth, 196 gorillas, 14 Gough Island (off Africa), 141 Grahame, Kenneth, The Wind in the Willows, 25 grass genomes, 75 gray wolf, 14 Great Barrier Reef, 76, 85, 153 Great Lakes, 144–45 green chemistry, 158 Green Revolution, 119 Greenland, 49 Greenpeace, 9–10 grizzly bear, 83 Gulf killifish, 85   hachimoji DNA and RNA, 105 Haemophilus influenzae, 66 Harbor View Park, Portland, Maine, 38 Haudenosaunee, 138 Hawaii: disease threat to endemic forest ¯ hi‘a lehua tree, 149 birds of, 146–47; ‘O Hayley Wood (UK), 2, 3, 13 heather and heathland, 78 Heck cows, 160–61, 162 Hendra virus, 145 Henslow, John, 53

INDEX 273

herbicides. See pesticides, herbicides, and insecticides heterozygotes, 62 Highland cattle, 41, 42 Himalayas, 39 homozygotes, 62 honeybees, 18–19, 127 Hooker, Joseph, 54 Hopkins, Gerard Manley, 1 horizontal gene transfer, 74–75, 125 horseshoe crab, 156 house sparrow, 78 howler monkey, 14 Hughes, Francine, 41 human environments: conservation management in, 37–40; genetic effects of, 77–80; hybridization due to, 85; natural and human landscapes, juxtaposition of, 1–4, 9, 13; naturalness, synthetic biology, and conservation issues in, 201–5, 211; nature seeking opportunities in, 37–40. See also agriculture Human Genome Project, 66, 117 humans: antibiotics, microbial resistance in, 80; composition of microbiome, 76; in eco synthetic biology, bodies as sites for, 118; genetic transfer from microbes, 75; hybridized ancestry of, 73–74; nature, human transformation of, 1–2, 6–9; organ transplants, cross-species, 111; zoonotic viruses, 145–46 hybridization, 70, 73–74, 77, 83–85, 137–38, 154, 162, 202, 209   Iberian ibex, 162 Iberian lynx, 82 iGEM (International Genetically Engineered Machine) competition, 113–14, 132 illegally traded species, 86 Impossible Foods, 108 in eco synthetic biology, 117–36; concerns regarding, 117–18, 123–25, 131–32, 134–35; for conservation purposes, 140; domestic crops and animals, 118–21; engineered gene drive, 129–33;

274

INDEX

in eco synthetic biology (cont.) gene flow/genetic transfer or introgression, 125–29; pesticides, herbicides, and insecticides, 121–23 inbreeding and low genetic diversity, 60, 68–69, 73, 82–83 India: Bt crops in, 122; decline of bird species in, 8; DIYbiosphere Project in, 113; GM crops in, 109, 122; naturalized plants in, 37; Oxitec mosquito releases, 130; reforestation efforts in, 34; regulation of synthetic biology in, 179; wolves in, 84 Indigenous peoples: American bison and, 208; American chestnut and, 136–38; Convention on Biological Diversity (CBD), protections in, 116; formal statements and declarations on genetic technologies, 183; giant anteaters and, 45; IUCN report and, 172, 173; rights and interests of, 7, 35, 48, 116, 173, 183, 189, 209 induced pluripotent stem cells, 162 Industrial Revolution, 3, 24, 112 insecticides. See pesticides, herbicides, and insecticides insulin, synthetic, 94 interleukin 2, 105 International Agency for Research on Cancer, World Health Organization, 126 International Genetically Engineered Machine (iGEM) competition, 113–14, 132 International Institute of Tropical Agriculture, Ibadan, Nigeria, 120 International Rice Research Institute, Philippines, 121 International Space Station, 4 International Treaty for Plant Genetic Resources for Food and Agriculture, 176 International Union for Conservation of Nature (IUCN): Bonn Challenge, 34; on conservation use of synthetic biology, 140, 172–73; on de-extinction, 164; list of invasive species, 38; Red List

of Threatened Species, 34, 71, 141; threat status and acceptability of genetic intervention, 187–88 invasive species, 29–32, 37–38, 86–87, 125,140–49, 153, 157, 198, 210 Ireland, human transformation of nature in, 35–36 isoprene, 159 IUCN. See International Union for Conservation of Nature   J. Craig Venter Institute, 89 jack pine warbler (Kirtland’s warbler), 30–31 Jaenisch, Rudolf, 94 jaguars, 47, 69 Japan: coral reefs, as climate refuge for, 36; Fukushima Daiichi nuclear disaster, 39; hachimoji DNA and RNA, 105 Japanese knotweed, 38 Japanese macaques, 39 Japanese/Korean chestnut, 136, 137 JCVI-syn1.0 (Synthia), 89 Johannsen, Wilhelm, 59 jumping genes/transposons, 75–76 Jungle Book (Kipling), 84 “junk” DNA, 75, 193 Jurassic Park (film), 49, 159, 160, 165 Jutland, Denmark, open heathland of, 78   Keats, John, 25 Kew Gardens (UK), 81 Kipling, Rudyard, Jungle Book, 84 Kirtland’s warbler (jack pine warbler), 30–31 Knight, Tom, 19 koalas, 87 Konik ponies, 41, 42 Korean/Japanese chestnut, 136, 137   laboratory mice and rats, 66, 142 Lagerfeld, Karl, 52 Laikipia District, Kenya, elephant conservation management in, 32–33 Lake District (UK), 25, 186 Lakenheath Fen (UK), 194–95



Lakota origin story, 209 LAL (limulus amebocyte lysate), 156 land reclamation, 157 landraces, 115 landscape genetics/genomics, 64–65 Last Universal Common Ancestor (LUCA), 49, 72, 106 leather, synthetic, 109 legal controls. See regulation of synthetic biology lemurs, 52 leopards, 14–15 Leopold, Aldo, 196, 212 Library of Congress, US, 104 limulus amebocyte lysate (LAL), 156 Linné (Linnaeus), Carl von, 45–46, 51 Linnean Society, 54 Liu, David, 100 Lloyds of London, 90 “localized” or “locally fixed allele” gene drives, 132, 149, 181 London, Jack, White Fang, 84 long-horned beetles, 145 LUCA (Last Universal Common Ancestor), 49, 72, 106 Lyell, Charles, 54 Lyme disease, 183 lynx, 41, 82   Maasai Mara, Africa, 4 MacArthur Foundation, x Mace, Georgina, xi, 171 maize. See corn/maize malaria, 130–32, 146–47, 171, 183, 199 mammoth, 109, 160, 162–63, 205 Man and Nature (Marsh), 24, 196 maned wolf, 47, 51 Maori, 183 marine environments: Convention on the Law of the Seas., 176; corals and coral reefs, 76, 85, 144, 153–55, 188; invasive/feral species in, 141–45; marine predators, 8; marine worms, 49; oceanic islands, 141–44. See also fish and fisheries Marsh, George Perkins, Man and Nature, 24, 196

INDEX 275

marsh harrier, 194 Masai giraffe, 71 Mascarene Islands, 38 Max Planck Institute of Molecular Plant Physiology, 110 Maynard, Charles, 138 McCartney, Stella, 109 meat, synthetic or cultured, 107–8 Memphis Meats, 108 Menangle virus, 145 Mendel, Gregor, 55–56, 90 Merkel, Hermann, 137 Mesteth, Wilmer, 209 methane reduction efforts, 158–59 methylation, 63, 64 Mexican wolf, 14 Mexico: Dia del Muerte in, 159; GM soy in, 127; monarch butterfly in, 128; screw-worm fly in, 93; whooping cranes in, 20 mice, 32, 66, 72, 141–43, 148, 183, 198 Michigan Department of Natural Resources, 31 microassisted migration, 155 microbiome: antibiotics, microbial resistance to, 80; biodiversity of, 15–16, 47, 49–50; conservation-based use of gene editing, 152–53; CRISPR technology, development of, 99–100; domestication of, 91; ecosystem microbiomes, 76; gene transfers within and from, 74–77; nitrogen-fixing microbes, as fertilizer, 120, 128. See also bacteria; viruses Microsilk, 109 Miescher, Friedrich, 56 migratory bird routes, 20–22 Millennium Seed Bank (Kew Gardens, UK), 81 Millett, Piers, 132 Mintz, Beatrice, 94 Modern Meadow, 109 monarch butterfly, 128 Mongolian wolf, 14 Monsanto, 9, 47, 122, 127, 139 Morton, Oliver, 90 Mosa Meat, 108

276

INDEX

mosaic cells, 63 mosquitoes and mosquito control, 17, 130–33, 146–47, 199 moths, 17–18, 93 Mount Saint Helens eruption, 192 mountain lions/panthers/pumas, 68–70, 72, 73, 82, 83, 87 Muir, John, 24 Murrill, William Alphonso, 137 mushrooms, 75, 178 mutation, 62–63   Nagoya-Kuala Lumpur Supplementary Protocol on Liability and Redress, 116, 176–77 nanotechnology, DNA, 104 National Environmental Policy Act (NEPA; US), 165 National Geographic, 160, 162 National Nature Reserves (UK), 3 National Trust (UK), 41–42 Native Americans, 136–38, 183, 208, 209 natural selection, 54, 61, 63 naturalized species, 37–38 naturalness, synthetic biology, and conservation, 191–212; adaptation, change, and evolution, 206–8; American bison, conservation of, 208–10; artificial versus natural, xi, 20–23, 42–44, 194–97, 207; biodiversity and genetic intervention, 191–94; challenges of, 211–12; concepts of nature and concerns about, 196–201; human-influenced ecosystems and, 201–5, 211; rewilding movement and, 206–8; salmon migration and, 191–93; technology, human ambivalence about, 196, 210–11 nature, 1–19; artificiality versus, xi, 20–23, 42–44, 194–97, 207; biodiversity at multiple levels of, 13–17 (See also biodiversity); boundless and endless, viewed as, 5–6; commercial/practical applications of synthetic biology, as resource for, 114–17; concept of, 23–27, 42–44, 199–201; conservation of, 1–4 (See also conservation management;

conservation using synthetic biology); domestic crops and animals, altering genomes of, 17; ecosystem, concept of, 13–14; equilibrium paradigm, breaking, 27–29; essentialist view of, 199; GM crops/organisms and concept of naturalness, 9–13, 17, 23; growing strangeness of, ix, x, 205; historical development of idea of, 24–25; human and natural landscapes, juxtaposition of, 1–4, 9, 13; human transformation of, 1–2, 6–9; in human-created environments, 37–40; past naturalness versus future naturalness, 39–40; pesticides and fertilizers, concern over effects of, 12; remaining wild lands, priority given to, 7–8; scientific names of species, table of, 213–20; “spaceship earth” model of, 4–6; species, concept of, 14–15; synthetic biology and conservation of, ix–xii, 17–19, 135–66 (See also conservation using synthetic biology); synthetic biology interacting with (See in eco synthetic biology); wild organisms, altering genomes of, 17–18 Neanderthals, 74 Neolithic (Agricultural) Revolution, 2, 3, 35–36, 53, 78 NEPA (National Environmental Policy Act, US), 165 New Guinea, 49 New York Botanical Gardens, 137 New York State Agricultural Experiment Station, 18 New York Zoological Society (now Wildlife Conservation Society), 208 New Zealand: invasive/feral species in, 143–44, 147; Maori, on genetic modification, 183; possum control in, 32; regulation of synthetic biology in, 179; tuatara conservation in, 72–73 Nipah virus, 145 nitrogen-fixing microbes, as fertilizer, 120, 128 nocturnality, 79 nonequilibrium ecology, 28–29 northern quoll, 87



Norway: farmed and wild salmon, gene flow between, 128; Svalbard Global Seed Vault, 81 novel ecosystems, 37, 43, 77 novel life-forms, 133–34 nuclear exclusion zones, 39   oak trees, 1, 49 Obama, Barack, 178 ocean. See marine environments Oglala Lakota, 209 ¯ hi‘a lehua tree, Hawaii, 149 ‘O Okavango Delta, Africa, 39 omega-3 fatty acids, 156 open access versus patents, 103, 116 orangefoot pimpleback, 51 orangutans, 14 organ transplants, cross-species, 111 organic farming and gene flow, 126 organochlorine pesticides, 12 Orpheus and Eurydice myth, 159 Oryx and Crake (Atwood), 167–69 oxalic acid, 138 Oxitec, 18, 130   Pakicetus, 77 Panama, Oxitec mosquito releases in, 130 Panamanian golden frog, 82 panthers/pumas/mountain lions, 68–70, 72, 73, 82, 83, 87 passenger pigeon, 14, 160, 212 Patagonia (outdoor clothing manufacturer), 109 Patent Act of 1952 (US), 116 patents: legal patentability of organisms, 115–16; open access versus, 103, 116 Patuxent Wildlife Research Center, Maryland, 21 Penicillium molds, 91 perfumes and cosmetics, 110 pesticides, herbicides, and insecticides: commercial/practical applications of synthetic biology in, 121–23; concern over chemical products, 12, 223n36; as conservation tool, 32; crops, engineering pest resistance in, 47, 121–22,

INDEX 277

126–27; engineered gene drives, 130–31, 148; gene flow/genetic transfer or introgression, 128, 129; genomes of pests, altering, 17–18; pests’ resistance to/tolerance of, 47, 79–80, 121, 122, 126–27; poison bait, 32, 141–42; sterile insect techniques, 92–93, 130, 131, 146–47 pharmaceuticals and “pharming,” 110–11 phenotype, 69 Philippines: anti-GM activism in, 120; International Rice Research Institute, 121; regulation of synthetic biology in, 179 Phoenix Zoo, Arabian oryx herd at, 82 photosynthesis, 79, 106, 120–21, 153 pigs, 111, 137, 158, 161, 167, 185, 198 pink bollworm, 93, 128 pink fairy armadillo, 49 Pivot Bio, 120 Plato, 70 Pleistocene rewilding project in Siberia, 40–41 pluripotent stem cells, 162 pollution cleanup, 157–59 poplar trees, 158–59 Post, Mark, 107–8 Powell, William, 138 prairie dogs, 34, 140 precautionary principle/approach, 182–83 Predator Free 2050 program, New Zealand, 143 predator species: conservation management asserting control over, 31, 32; marine predators, 8; rewilding movement and reintroduction of, 40, 41 product substitution, 155–57 prokaryotes, 57, 58 Prometheus myth, 196 protection model of conservation, 1–4 PROVEN, 120 Pseudomonas syringae, 123 psilocybin, 75 public health: applications of genetic technologies for, 183; Chernobyl disaster, threat from, 186–87

278

INDEX

public trust of/engagement in genetic technologies, 183–87 public and popular involvement in synthetic biology, 113–14 pumas/panthers/mountain lions, 68–70, 72, 73, 82, 83, 87 purple hairstreak butterfly, 1 Pyrenean ibex, 162 Pythagoras, 53   quagga, 14, 161   rabies, 34 raccoon dog, 39 radiation: Chernobyl, nuclear fallout from, 186–87; deliberately introducing mutations through, 92–93, 137 rainbow trout, 108 rats, 32, 141–44, 148 Ray’s Flora of Cambridgeshire, 1 recombinant DNA technologies, 93–96, 156 red fox, 143, 202 Redford, Kent, x, xi, xii, 4, 38, 45, 46, 136, 140, 172–73, 203 Registry of Standard Biological Parts, 103 regulation of synthetic biology: bio­ technology industry, role of, 175; de-extinction issues, 165; democratic process and, 175; different applications, treatment of, 180–82; digital DNA sequencing, 177; engineered gene drives, 132, 181; gene editing tech­ nologies, 179, 181–82; GM crops and organisms, 178–79, 181; governments’ role in, 175–76, 177–80; Indigenous peoples, rights and interests of, 183; international conventions on, 176–77; IUCN report, criticism of, 172–73; patentability of organisms, 115–16; product-based versus process-based approach to, 179; public trust and engagement, 183–87; US National Academies classification of biotechnology products, 170–71. See also specific Acts and Conventions reindeer, 186

restoration of ecosystems, as conservation goal, 34–35, 36, 41, 201–2 reversal gene drives, 132, 148, 181 Revive & Restore, 160, 163 rewilding movement, 40–42, 206–8 rhea, 47 rhino horn, synthetic, 156–57 rice, 53, 68, 91, 95, 110, 119–21, 126 Rio Declaration on Environment and Development, 182 Riyadh, Saudi Arabia, Arabian oryx herd at, 82 RNA (ribonucleic acid), 57–59, 63, 105, 142, 231–32n26 RNA editing, 104 RNA silencing/RNA interference (RNAi), 122–23, 145, 159 robin, American versus European, 52 Rockefeller Foundation, 119 Rodrigues fody, 38 Romanticism and nature, 24 Romesberg, Floyd, 105 Roosevelt, Teddy, 208–9 Roslin Institute, Scotland, 161 rough fatmucket, 51 Roundup and Roundup Ready crops, 9, 47, 126–27 Royal Society (UK), 89–90, 179, 185 Royal Society for the Protection of Birds (RSPB), 194 Royal Society Te Apārangi (New Zealand), 144   sad elliptio, 51 salamanders, 150 salmon, 83, 96, 108, 128–29, 191–93 San Diego Zoo, 67, 81, 162 A Sand County Almanac (Leopold), 196 Sarloos wolfdog, 160 scents and cosmetics, 110 Schwille, Petra, 106 scientific nomenclature for species: historical development of, 51–52, 71; table of scientific names of species, 213–20 SCIMAC (Supply Chain Initiative for Modified Agricultural Crops), 10 ScottsMiracle-Gro, 127



screw-worm fly, 93 Scripps Research, 105 sea lamprey, 144–45 selective breeding, 90–93, 95, 115, 160–61 Selous, Africa, 4 SER (Society for Ecological Restoration), 40 Serengeti, Africa, 4 sexual reproduction, 61–62 Shaw, Pamela, 38 Shawver, Laura, 105 sheep, 14, 52, 82–83, 90, 158, 161, 165, 186 Shelley, Mary, Frankenstein, 12–13, 159, 160, 167–69, 196 Siberia, Pleistocene rewilding project in, 40–41 Silent Spring (Carson), 12, 43, 196 silk, 109 Sina Bear Eagle, 209 Sites of Specific Scientific Interest (UK), 3 “smart” bacteria, 152 Smokejumper 3 (hot spring, Yellowstone), 15 Snuppy the dog, 161 Society for Ecological Restoration (SER), 35, 40 Society for the Preservation of the Wild Fauna of the Empire (now Fauna and Flora International), 14 Society for the Promotion of Nature Reserves (UK), 2–3 soil nematodes, 66 somatostatin, 94 sooty terns, 148 The Sorcerer’s Apprentice (film), 160 South Georgia Heritage Trust, 32 South Georgia pipit, 32 soy leghemoglobin heme protein, 108 soy/soya/soybeans, 8, 11, 47, 95, 108–10, 126–27 “spaceship earth” model of nature, 4–6 sparrows, 78 species: biodiversity of, 14–15, 16, 47, 50; classification issues, 68–74; concept of, 14–15, 16; conservation organizations building legislation around, 71–72; conservation-reliant, 31; de-extinction

INDEX 279

critique and, 163–64; DNA “barcodes,” species identification via, 86; Endangered Species Act (US) and, 31; endemic, 47, 48; extinction of, 49, 50; feral/invasive, 32, 37–38; genes crossing between, 74–77; genetic variation and speciation, 61–65; hybridization of, 70, 73–74; IUCN Red List of Threatened Species, 34; as natural unit versus human-designated category, 70–72, 73; only surviving members of now-extinct taxa, 72–73. See also scientific nomenclature; specific species by common name species action plans, in UK, 72 spiders/spider silk, 52, 109, 114 Sputnik I, 5 squalane, 110 sterile insect techniques, 92–93, 130, 131, 146–47 Stevenson, Adlai, 5 stoats, 143 “straying” salmon, 192 Streisand, Barbra, 161 student/popular involvement in synthetic biology, 113–14 subspecies, 14–15, 69–70 Supply Chain Initiative for Modified Agricultural Crops (SCIMAC), 10 sustainable development movement, 24 Svalbard Global Seed Vault, Norway, 81 Sweden, European vipers in, 83 sylvatic plague, 34, 140 Syngenta, 120, 123 synthetic biology, 89–106; animal-­ machine hybrids, 104–5; “cell-free,” 103–4; creation of first synthetic cell, 89, 96; data-storage unit, using DNA as, 104; defined, 89–90, 96–97; designbuild-test-learn cycle in, 101–2; DNA nanotechnology, 104; domestic organisms, selective breeding of, 90–93, 95; efficiency requirements, automation, and acceleration, 102–3; gene editing technologies, 96–101 (See also gene editing technologies); genome editing, 99–100, 101, 108; GMOs and, 95; hachimoji DNA and RNA, 105; natural

280

INDEX

synthetic biology (cont.) world, interacting with (See in eco synthetic biology); patents versus open access in, 103; radiation and chemicals, deliberately introducing mutations through, 92–93, 137; recombinant DNA technologies, 93–96; RNA editing, 104; technological system, treating biology as, 97–98; xenobiology, 105, 134. See also commercial/ practical applications of synthetic biology; conservation using synthetic biology; debates about synthetic biology; naturalness, synthetic biology, and conservation synthetic ecology, 152–53 Synthia (JCVI-syn1.0), 89   TALENs, 99 Target Malaria, 130–31 Tasmanian devil, 82, 145 Tasmanian tiger (thylacine), 164 taxonomic classification of species, 68–74 Tears of the Trufflepig (Flores), 169 technology, human ambivalence about, 196, 210–11 termites, 45–48 terraforming, 134 Texas panther, 69–70, 83 textiles, 109 thylacine (Tasmanian tiger), 164 tilapia, 108 tomatoes, 12, 92, 108–9, 115, 120 transgenesis, 95–96, 120, 138–39 transplanted organs, cross-species, 111 transposons/jumping genes, 75–76 trees and forests: ancient woodlands, 1–4, 9; Atlantic Forest, Brazil, 4; as carbon sinks, 159; church forests, Ethiopia, 4; deforestation, 35–36, 47; rain forests, 3, 13, 14, 25, 46, 47, 160; reforestation, commitments to, 34; US Forest Service, 31. See also Hawaii; specific types of tree Tristan albatross, 141

“Trojan horse” fears, 139, 171, 249n10, 257n8 trophic cascades, 148 Trump, Donald, 178 Tschermak, Erich von, 56 tsetse fly, 93 Tu Youyou, 110 tuatara, 72–73 tube worms, 49   ultralight aircraft, assisting birds on migratory routes with, 20–22 United Kingdom (UK): Australia, introduction of invasive/feral species into, 143; cranes in, 194–95; Dutch elm disease in, 61; European wildcats in, 84; GM crops and organisms in, 9–11; nature and conservation, development of concepts of, 3, 25; species action plans in, 72; urban environments, red foxes adapting to, 202. See also specific locations, organizations, agencies, and Acts United Nations (UN): Convention on Biological Diversity (CBD), 16, 35, 72, 116, 140, 176–77; Convention on International Trade in Endangered Species (CITES), 72, 176; Convention on the Law of the Seas, 176 United Nations Environment Programme (UNEP), 140 United States (US): Bt crops in, 122; “command and control” approach to conservation management in, 29–30; DIYbiosphere Project in, 113; GM crops and organisms in, 11, 109; nature and conservation, development of concepts of, 24–25; Oxitec mosquito release plans, 130; regulation of synthetic biology in, 178; urban environments, coyotes adapting to, 202. See also specific locations, organizations, agencies, and Acts urban ecosystems, 37, 78–79, 202 US Department of Agriculture (USDA), 178



US National Academies, 112–13, 154, 170, 178, 188 utilitarian conservation movement, 24   VanderMeer, Jeff, Annihilation, 168 Venter, Craig, 66, 89, 96, 104 viruses, 9, 15–16, 18–19, 47, 50, 57, 66–67, 75, 93–94, 99–100, 111, 115, 122–23, 145–46, 153   Walking with Dinosaurs (TV show), 49 Wallace, Alfred, 52–55; “On the Tendency of Species to Form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection” (with Darwin), 54; “On the Tendency of Varieties to Depart Indefinitely from the Original Type,” 53 wasps, invasive, in New Zealand, 143–44, 147 Watson, James, 7, 56 Weaver, Sigourney, 165 West Nile virus, 145 western corn rootworm, 123 whales, 77 wheat, 2, 53, 91, 119, 123, 126, 138, 204 White Fang (London), 84 White Oak Plantation, Florida, 68–69 white-nose syndrome, 149 white-tailed deer, 203 whooping cranes, 20–22, 195 Wicken Fen National Nature Reserve (UK), 41–42 Widestrike 3, 122

INDEX 281

Wilderness Act of 1964 (US), 29, 200 Wildlife Conservation Society (formerly New York Zoological Society), 208 Wind Cave National Park, SD, 208–9 The Wind in the Willows (Grahame), 25 The Windup Girl (Bacigalupi), 169 Winkler, Hans, 65 Wolbachia, 146–47 wolves, 14, 15, 34, 39, 41, 47, 51, 84, 90, 160 Wood Buffalo National Park, Canada, 20 woodlands. See trees and forests woolly mammoth, 109, 160, 162–63, 205 World Conservation Congress, 172 World Economic Forum, 112, 113, 114, 116 World Health Organization, 126 Wyalusing State Park, WI, 212   xenobiology, 105, 134 XNA (xeno-nucleic acid), 134 X-rays, 92   yeast, 66, 108, 109, 110, 114 Yellowstone National Park (US), 4, 15, 41, 229n58 Yosemite (US), 4, 24 Yurok Tribal Council of the Klamath River, 183   Zahniser, Howard, 200 Zika, 130 zinc fingers, 99 zoonotic viruses, 145–46